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The CANADIAN 
FIELD-NATURALIST 



/ 



Published by THE OTTAWA FIELD-NATURALISTS' CLUB, Ottawa, Canada 








Volume 116, Number 1 



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■^^s^/p-v/u^fGjO 




Januarv-March 2002 



The Ottawa Field-Naturalists' Club 

FOUNDED IN 1879 

Patrons 

Her Excellency The Right Honourable Adrienne Clarkson, C.C, C.M.M., CD. 

Governor General of Canada 
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Cover: Wood Bison, Bison, bison bison photographed at Wood Buffalo National Park. Photo courtesy W.A. Fuller. See: 
Canada and the "buffalo" pages 141-159. 



THE CANADIAN 
FIELD-NATURALIST 



Volume 116 
2002 



THE OTTAWA FIELD-NATURALISTS' CLUB 

Ottawa Canada 



The Canadian Field-Naturalist 



Volume 116, Number 1 



January-March 2002 



r 
I 



Spread and Disappearance of the Greater Prairie-Chicken, 
Tympanuchus cupido, on the Canadian Prairies and adjacent areas 

C. Stuart Houston^ 



'863 University Drive, Saskatoon, Saskatchewan S7N 0J8 Canada 

Houston, C. Stuart. 2002. Spread and disappearance of the Greater Prairie-Chicken, Tympanuchus cupido, on the 
Canadian prairies and adjacent areas. Canadian Field-Naturahst 116(1): 1-21. 

In western Canada, the Greater Prairie-Chicken (Tympanuchus cupido) has come and gone. Following closely on agricul- 
tural settlement, thriving on grain as an alternate food, the "Pinnated Grouse" reached Winnipeg in 1881 and Carberry, 
Manitoba, in 1886; Indian Head, Saskatchewan, in 1895; and Hanna, Alberta, in 1911. It spread as far northwest as Lac la 
Biche, Alberta. Once half the land in a given area was broken, this species diminished in numbers, retreating to local areas 
of thick grass around sloughs and lakes. By the late 1930s almost all were gone. Habitat factors such as fragmentation and 
separation of grasslands were further accentuated by cattle overgrazing, burning, and drought, and then by hybridization of 
surviving, isolated birds. By accessing records for the prairie provinces from nine unpublished or not-readily-available 
sources (unavoidably skewed by a preponderance of Saskatchewan records), I offer documentation of increases and 
decreases, as well as an assessment of the species' status in northeastern Montana and northwestern Ontario. I have also 
catalogued 131 Northern Great Plains egg sets from 36 North American oology collections. Together this material allows a 
more complete assessment than has been possible previously. 

Key Words: Greater Prairie-Chicken, Tympanuchus cupido, spread, decline, extirpation, ecologic factors. 



On the Canadian grasslands, the Greater Prairie- 
Chicken (Tympanuchus cupido) has come and gone. 
Because there is concern for many other species of 
grassland birds, including the Sharp-tailed Grouse 
(Tympanuchus phasianellus), and because two other 
species, the Sage Grouse (Centrocercus urophasi- 
anus) and Burrowing Owl (Athene cunicularia), may 
be facing extirpation in Saskatchewan, it seems 
worthwhile to summarize the spread and demise of 
the Greater Prairie-Chicken in western Canada and 
adjacent North Dakota. Are there common threads or 
lessons to be learned? 

In this review I bring together, before it is too late, 
published and previously unpublished information. 
To provide localities and dates, and especially breed- 
ing records, I made particular use of: (a) the Andrew 
Graham/Thomas Hutchins combined report from 
Hudson Bay, 1772, (b) David Douglas' Manitoba 
observation of 1827; (c) responses to Mrs. Isabel 
Priestly' s request (1943) for Saskatchewan records; 
(d) unpublished responses to my request for addi- 
tional Saskatchewan records in 1956; (e) George J. 
Mitchell's 1959 Alberta's Upland Game Bird 
Resource (1959); (f) Margaret Belcher's 1961 report 
of museum records in Saskatchewan; (g) Dale 
Hjertaas' 1988 summary of Saskatchewan records 
since 1966; (h) Anita Steeg Kovacs' 1984 summary 
of newspaper accounts in southern Manitoba, 



1884-1949, (i) a Montana sportsman's magazine 
article that adds extreme northwest Montana to their 
maximum range, (j) Harry Lumsden's previously 
unpublished reports that add northwestern Ontario to 
their range, and (k) oology records for the three 
prairie provinces and the Dakotas in major museums. 
This information has been brought together in one 
place for use by future researchers, even though 
most of the Saskatchewan locations have been 
mapped in Smith (1996). The dearth of readily avail- 
able accounts largely explains the paucity of western 
Canadian information in Schroeder and Robb 
(1993). 

Problem of name 

The names used for prairie grouse could hardly be 
more confusing. Sadly, "Prairie Chicken" was, and 
still is, the name almost universally applied to the 
Sharp-tailed Grouse through western Canada. Ernest 
E. Thompson [Seton] in his 1890 account of the 
Birds of Manitoba, used "Prairie Chicken" for his 
account of the Sharp-tailed Grouse, and used "Prairie 
Hen" or Pinnated Grouse for what was then known 
to ornithologists south of the boundary as the Greater 
Prairie-Chicken. Seton's usage followed that of 
Macoun (1883). Macoun and Seton thus elevated the 
use of "Prairie Chicken" for the Sharp-lailcd Grouse 
far beyond that of a simple and misleading folk 



The Canadian Field-Naturalist 



Vol. 116 



name and gave such use scientific standing for more 
than half a century. The general public and even 
game guardians and museum employees used 
"Prairie Chicken" much more often for the Sharp- 
tailed Grouse than for the Greater Prairie-Chicken. 

For this reason, most popular accounts that refer 
to "prairie chickens" are suspect. In fact, nearly all 
of them refer to the Sharp-tailed Grouse. Johnston 
and Smoliak's helpful review of the Greater Prairie- 
Chicken in 1976 fell into this trap when they gave 
credence to Bulyea (1901) who unquestionably 
referred to Sharptails between Moosomin and 
Qu'Appelle, Saskatchewan; they might better have 
quoted Rutherford (1914), who did recognize the 
presence of Greater Prairie-Chicken, and, differenti- 
ated between the two species. To avoid confusion, 
Cooke (1885), Roberts (1936) and Johnsgard (1973) 
used the term "Pinnated Grouse." Although 
"Pinnated Grouse" referred to only one subspecies of 
the Greater Prairie-Chicken, and although common 
names are no longer assigned to subspecies, 
"Pinnated" is the only term that unequivocally indi- 
cates that past observers, at a given time or place, 
saw the Greater Prairie-Chicken. 

Problem of identification 

In an ideal world, one would anticipate few prob- 
lems of mistaken identification between two grouse 
species that show such marked differences, one from 
the other. First, those reports that specifically men- 
tioned the dark, rounded or squared tail, and the 
heavily barred underparts, offer proof of the Greater 
Prairie-Chicken, in sharp contradistinction to the 
light-coloured, elongated 'pin-tail' and finer v- 
shaped breast markings of the Sharp-tailed Grouse. 
Although the Ruffed Grouse {Bonasa umbellus) also 
has a rounded tail, its restriction to wooded areas 
should exclude it from consideration in most plains 
localities. Second, the mating rituals at the "booming 
grounds" of the Greater Prairie-Chicken and the 
"dancing grounds" of the Sharp-tailed Grouse are 
different. Third, as a game bird, individuals from 
each fall kill were available for close inspection, pro- 
viding in-the-hand identification opportunities not 
available for many non-game species. The most 
credible observers are those who mentioned the two 
species together, for these folks clearly had an 
opportunity for comparison. Nevertheless, one 
would be naive to expect uniform accuracy, because 
wishful thinking, a fertile imagination, or a faulty 
memory may on occasion have contributed to occa- 
sional and innocent misidentifications, particularly 
of sightings since 1940 — and we know not which 
or how many of these have been reproduced here. 

Initial range 

It is somewhat ironic that the Greater Prairie- 
Chicken was becoming extirpated in the eastern 



United States even as it spread west across the 
plains. For example, it disappeared from Tennessee 
in 1850, Kentucky in 1874, Arkansas in 1913, and 
Ohio in 1934 (Schroeder and Robb 1993), and its 
decline in northern Illinois began as early as 1850 
(Schorger 1944). "At one time it was the leading 
upland game bird of the grasslands of central North 
America" (Robel et al. 1970). In Wisconsin in 1850, 
"Two sportsmen, with one dog, generally bag from 
fifty to eighty in a day" (Hoy 1852). "Thousands of 
barrels of them" were sold by market hunters (Rue 
and Allen 1973). 

At the time of the return trip of the Lewis and 
Clark expedition in 1806, Greater Prairie-Chickens 
were present in Illinois. The sighting "highest up" 
the Missouri was at the James River, (now Yankton, 
South Dakota), on 2 September 1806 (Burroughs 
1961). 

T. S. Roberts (1936) stated "There were no 
Pinnated Grouse in Minnesota in the days of the 
early explorers. ... [this species] extended its range 
westward and northwestward just as the country was 
settled, keeping pace with the forefront of grain- 
fields." Through 1850 to perhaps 1880, the range of 
overlap between the Pinnated Grouse and the Sharp- 
tailed Grouse range was very narrow, because the 
one species ceased where the other began (Johnsgard 
and Wood 1968). 

The IS'^'-century record for Ontario 

Previous accounts have overlooked Pennant's 
1792 statement, "Is frequent about a hundred miles 
up Albany river, in Hudson's Bay." This statement is 
apparently derived from the Andrew Graham unpub- 
lished manuscript E.2/9, Hudson's Bay Archives, 
Provincial Archives of Manitoba, written probably in 
1772: "Pinnated Grous [sic] is found about Henley 
Settlement in Hudson's Bay. Legs covered with soft 
brown feathers, toes naked and pectinated. The tufts 
which distinguish this species from all others are 
rooted high on the neck, not far from the hindpart of 
the head." (Houston et al. 2002). Henley House was 
situated just below the junction with the Kenogami 
River, about 160 km upstream from Albany on 
Hudson's Bay. 

In 1828, the farthest north known record in 
Ontario of the Greater Prairie-Chicken was in Essex 
County, about 1000 km south of Henley House. As 
settlers cleared land, it spread eastward to Toronto 
by 1858 and north to Holland Marsh at the south end 
of Lake Simcoe by 1875. It then retreated, and was 
extirpated from the province by 1924 (Lumsden 
1966). Next, the species spread north in Michigan to 
Sault-Ste. Marie, then gradually colonized 
Manitoulin Island, from west to east, between 1941 
and 1962, before being swamped through hybridiza- 
tion with the Sharp-tailed Grouse, newly arrived 
from the north (Peck and James 1983). 



2002 



Houston: Greater Prairie-Chicken on the Canadian Prairies 



I 
I 



Overlooked record near present Winnipeg 

All previous accounts, save that of Roberts 
(1936), have overlooked an important record. In the 
first six days of August 1827, David Douglas, under 
the heading Tetrao cupido, "killed several birds of 
this species between Red River and Pembina in 49° 
north latitude. This may, perhaps, be found to be its 
most northern range" (Douglas 1829). Douglas 
arrived at Fort Garry, within present Winnipeg, on 
12 July 1827 and stayed until 10 August. He made a 
number of one-day botanizing trips and one two-day 
trip on 23-24 July from Fort Garry to Whitehorse 
Plain, 29 km west up the Assiniboine River. On 1 
August he went on horseback to a low hill about 27 
km east of Fort Garry (Douglas 1914). His Greater 
Prairie-Chickens must have been collected much 
closer to Fort Garry than to Pembina. In view of 
Douglas' single record of this species within 
Manitoba in 1827, Roberts' claim that there were 
none in Minnesota through 1850 is probably an 
error. 

Spread into Northwestern Ontario (Figure 1) 
The only published observation in northwestern 
Ontario, of "occurrence" with "no indication of 
breeding," is in an eight-page mimeographed list, 
Birds of the Canadian lakehead area, by K. Denis 
(1940, cited by Peck and James 1983: 137). 
Nevertheless, Greater Prairie-Chickens were present 
between 1 920 and 1 940 near Warroad, Minnesota, at 
the southwestern comer of Lake of the Woods, and 
east along the Rainy River, the United States- 
Canada boundary, to Fort Frances (The last booming 
was heard 5 km southeast of Warroad in 1963; J. L. 
Ruos, Leader, Grouse Research Project, Minnesota 
Game and Fish, letter to Harry G. Lumsden, 16 
December 1963). It also occurred within northern 
Minnesota as far east as Duluth until 1952 (Green 
and Janssen 1975). Slightly farther north, near 
Dryden, Ontario, Chief Ranger John Anderson in 
1944 reported to Lumsden that he saw 10 to 15 
grouse with yellow-orange neck patches displaying 
near the four-mile comer of the Rice Lake loop road, 
»east of Quibell. Ontario Conservation Officer A. R. 
lOlsen (six-page report to Lumsden, 1959) inter- 
jviewed long-time farmers west of Dryden and deter- 
[mined that, after they cleared the bush for farming, 
Jharp-tailed Grouse moved in and increased until the 
.920s, accompanied by a few Greater Prairie- 
'hicken, which were preferred for food because of 
leir larger size. On 8 April 1959, 5 km north and 1 
:m east of Minnitaki post office west of Dryden. 
[Olsen observed a dancing ground of 2 hectares on a 
stubble field with six birds. When flushed, two were 
positively identified as Pinnated Grouse and the 
other four as Sharptails. The above observations 
appear to add a new species to at least the hypotheti- 
cal bird list for Northwestern Ontario. 



Spread into Manitoba (Figure 1 ) 

The spread of the Greater Prairie-Chicken into, 
and across, Minnesota, beginning about 1850, has 
been well documented by Roberts (1936). Cooke 
(1885) reported from Moorhead, Minnesota, its inva- 
sion of the Red River valley of Manitoba: "Its north- 
ern range has always fallen short of our northern 
boundary until last summer, when they invaded 
Manitoba. Mr. C. W. Nash of Portage la Prairie, 
says: 'This autumn we had a curious influx of the 
Pinnated Grouse. I imagine that they, like some other 
birds, are following up civilization, for until last year 
(1884) none were seen by the Indians and half-breed 
hunters.'" 

The Greater Prairie-Chicken then spread west 
roughly in concert with white settlement. In 
Manitoba it reached Winnipeg in 1881, Portage la 
Prairie in 1882 (Nash in Cooke 1888), Carberry in 
1886 (Thompson 1890), and Aweme/Treesbank in 
1888 (Criddle 1929). Alex McArthur gave a talk on 
winter birds in January 1887 and reported that the 
previously absent Pinnated Grouse was "now quite 
common ... The reason for its rather sudden appear- 
ance and rapid increase seems to be that it has fol- 
lowed the wheat fields." C. P. Forge collected at 
least 17 Greater Prairie-Chicken egg sets near 
Carman, Manitoba, 1900-1904 (Table 1, Houston 
andBechard 1987b). 

Four decades after their arrival in Manitoba, they 
were still at a booming ground near Carberry 
(Symons 1967). They spread north as far as Swan 
River (specimen in 1938; Minish 1987a), Peonan 
Point (specimen) and Grand Rapids (sight record) 
(Godfrey 1966). According to T. Schindler. they 
reached their highest numbers at Oak Hammock 
Marsh in the 1920s (Gardner 1981). 

Spread into Saskatchewan (Figure 2) 

In Saskatchewan, Greater Prairie-Chickens spread 
and multiplied rapidly. One was killed at Indian 
Head late in 1895 (Macoun 1900). The first 
Saskatchewan nest was found at Crescent Lake, near 
Saltcoats, in 1897 (Table 1), where from 1910 to 
1920 they were as common as the Sharp-tailed 
Grouse (Baines 1956), a claim made also by Ferry 
(1910) at the Quill Lakes in 1909. Greater Prairie- 
Chickens reached Tullis after 1906 (Roy 1996). 
Their presence on intensively farmed land near 
Regina was short-lived, but they were numerous 
5 km west of that city on 30 September 1913 (H. H. 
Mitchell, in Belcher 1961), and were also seen 8 km 
south of that city on 27 April 1913 (Buchanan 1914). 
They reached the Battleford district in 1913. and 
there were "large flocks" at Lloydminster in 1914 
(Bradshaw 1915). In 1914. J. D. Soper (1970) found 
"a few" at a ranch 10 km south of Broadview, but 
when he returned from late April to early June 1921, 
he saw none, though residents reported a few still 



The Canadian Field-Naturalist 



Vol. 116 




Figure 1 . Localities mentioned in Manitoba. 



present. In 1915 George Lang reported large flocks 
of prairie grouse at Indian Head, approximately one- 
half being Greater Prairie-Chicken or hybrids (the 
only mention of hybridization located in the 
Saskatchewan game reports). At Indian Head in the 
winter of 19 16-1917 they were sufficiently common 



that George Lang, voluntary game guardian, 
observed seven dead birds within one mile; they had 
been killed by flying into telephone wires (Bradshaw 
1917). They appeared at grain baits at Raymore in 
the winter of 1916-17 with flocks of Sharp-tailed 
Grouse (Charles Harris in Wayne C. Harris, unpub- 



2002 



Houston: Greater Prairie-Chicken on the Canadian Prairies 




Figure 2. Localities mentioned in Saskatchewan. 



lished inanuscripts). In 1918, H. F. Ward at Pennock 
near Saltcoats reported a "surprising number" of 
Pinnated Grouse, including one covey of eleven, up 
to six or seven in other flocks, and several singles 
(Bradshaw 1919). 



At Ituna they were first seen about 1910, five or 
six years after initial settlement, when little land was 
yet broken; H. M. Rayner saw an occasional covey 
until the fall of 1943 when he flushed eight birds 
3 km NNE of Ituna (letter. 4 March 1956)^ Fred G. 




Figure 3. Localities mentioned in Alberta. 



Bard found them still common, evidently breeding, 
at Dafoe in both 1928 and 1929, and collected an 
adult specimen on 25 April in the latter year 
(Belcher 1961). 

The Greater Prairie-Chicken reached the northern 
limit of its range in longitude 102° at Kamsack in 
1907 (Fraser 1961). Farther west it reached Edam in 



1913 where W. E. Lake collected one on 13 
September (Belcher 1961). By 1914 it had reached 
the Onion Lake Indian Reserve, near the Alberta 
boundary north of Lloydminster (Bradshaw 1915). 
The species was already present at Somme when 
Wallace Black homesteaded in 1920, and it reached 
Tisdale in 1923 (Houston and Street 1959). By 1924 



2002 



Houston: Greater Prairie-Chicken on the Canadian Prairies 



Table 1. Greater Prairie-Chicken egg sets. Northern Great Plains (see acknowledgments for institute initials) 



# sets sets eggs/set 

ceggs 



Locality 



Day/Mo 



Year 



#eggs 



Collector 



Institution 



Unknown 

unknown 
unknown 
unknown 
unknown 



3 10.67 



20 May 



1905 



1930 



7 


J Kowal 


pma 


12 


J Kowal 


PMA 


(1) 


J Kowal 


PMA 


13 




ASC 



Alberta (AB) 


2 


2 


11.50 




23 






Stony Plain 






— 


1901 


14 


SSS Stansell? 


PMA 


Heatherdown 






— 


— 


9 


M PollackAVilby 


PMA 



Saskatchewan (SK) 

Crescent L 

Saltcoats 

Saltcoats 

Melville 

Wauchope 

Wauchope 

North Portal 

S of Gainsborough 

Wauchope 

5 mi nw Antler ND* 

5 mi nw Antler ND* 

4 mi nw Antler ND* 

5 mi nw Antler ND* 



13 



10.88 



24 May 



30 May 
20 May 
10 May 
3 May 
12 May 
25 May 
6 June 
29 May 
8 June 
14 June 



1897 
1915 
1915 
1920 
1923 
1924 
1925 
1925 
1925 
1925 
1927 
1927 
1927 



87 
11 

(2) 
10 
(4) 

(1) 
12 

14 

(1) 

11 

9 

10 

10 



Note: those four with ND* probably were in Saskatchewan 



F Baines 
N Jowsey 
N Jowsey 
HC Grose 
HH Pittman 
HH Pittman 
GF Abbey 
GF Abbey 
HH Pittman 
GL Davy 
GA Withey 
GL Davy 
GA Withey 



WFVZ 

PMA 

PMA 

SMNH 

USask 

SMNH 

PMYU 

RBWWF 

USask 

WFVZ 

WFVZ 

WFVZ 

SBCM 



Manitoba (MB) 

Shoal Lake 

nr Reabum 

Morris 

Beulah 

Long L nr Reabum 

Carman 

Carman 

nr Roland 

Carman 

Bamsley 

Salterville 

Salterville 

Myrtle 

Salterville 

Salterville 

Myrtle 

nr Roland 

Myrtle 

Myrtle 

Myrtle 

MadfordAVadhope? 

Myrtle 

Elm Point 

e of Carman 

Salterville 

MacDonald 

Lake Winnipeg 

Douglas 

Kalevala 

Kalevala 

Oaklands 

St. Francis Xavier 



32 



30 



10.03 



8 June 

9 June 
17 May 

29 Apr 

10 June 

30 May 

25 May 

28 May 

29 May 

30 May 

10 June 
15 May 
17 May 
17 May 

19 May 

31 May 
31 May 
21 June 

21 June 
2 July 

5 June 

20 June 
24 May 
24 May 

22 June 
2 June 

1 June 
28 May 
30 May 

1 1 June 
4 June 



301 

1894 
1894 
1896 
1899 
1899 
1899 

1900 
1901 
1901 
1901 
1901 
1902 
1902 
1902 
1902 
1902 
1902 
1902 
1902 
1902 
1903 
1903 
1904 
1904 
1905 
1905 
1916 
1921 
1921 
1927 
1928 



6 ne 
12 
? 

2 ne 
8 
13 



9 

13 

8 

11 
9ne 

8 

11 

8 

10 

17 

10 

15 

13 
10 ne 

10 

14 

9 

6 

10 

** 

12 
13 
6 
12 



E Arnold 

W Raine 

PB Peabody 

AJ Dennis 

W Raine 

JW Preston 

JW Preston 

CP Forge 

CP Forge 

WC Bradbury 

CP Forge 

CP Forge 

CP Forge 

CP Forge 

CP Forge 

CP Forge 

CP Forge 

CP Forge 

CP Forge 

CP Forge 

J Labarthe 

GE Congdon 

CP Forge 

CP Forge 

CP Forge 

CP Forge?AVilby 

DWilby 

CH Young 

ES Norman 

ES Norman 

AG Lawrence 

AG Lawrence 



I 



WFVZ 

NMC 

MPM 

DMW 

WFVZ 

ROM 

OSUM 

WFVZ 

OSUM 

DM 

NYSM 

WFVZ 

WFVZ 

WFVZ 

WFVZ 

MVZ 

WFVZ 

FMG 

WFVZ 

WFVZ 

UNR 

WFVZ 

WFVZ 

AMNH 

USNM 

PMA 

NMC 

NMC 

SBCM 

UMMZ 

NMC 

NMC 

Contitnu'ii 



The Canadian Field-Naturalist 



Vol. 116 



Table L Continued 



# sets 



Locality 



sets 
ceggs 



eggs/set 



Day/Mo 



Year 



#eggs 



Collector 



Institution 



North Dakota (ND) 52 

Wahpeton 

Church's Ferry 

Devils Lake 

Rock Lake, Towner Co. 

Pembina 

Pierce Co. [Buffalo Lake] 

Stump Lake 

Eddy Co. 

Eddy Co. 

Duck Lake 

Lakota 

Arthur, Cass Co. 

Rock Lake 

Ward Co. 

8 mi W Deering 

2 mi NW Antler 

4 mi W Deering 

nr Deering 

12miW Antler 

6 mi S Flaxton 
10 mi E Deering 

4 mi N Denbigh 

5 mi E Deering 

4 mi E Deering 

5 mi E Deering 

7 mi NE Flaxton 

2 mi E Ander 

3 mi NE Deering 

6 mi SW Antler 
N of Antler 

6 mi E Deering 

2 mi SW Antler 
5 mi SE Antler 

4 mi SE Antler 
Antler 

NE Deering 
NE Deering 
NE Deering 
NE Deering 
McHenry Co. 
McHenry Co. 
McHenry Co. 

3 mi SE Antler 
McHenry Co. 
McHenry Co. 
McHenry Co. 
McHenry Co. 
McHenry Co. 
McHenry Co. 
McHenry Co. 
McHenry Co. 
2 mi E Loraine 



50 



11.86 



South Dakota (SD) 
Yellowstone R. 
Vermilion 
Vermillion 
Vermillion 



28 



26 



10.96 







593 






— 


— 


11 


JK Jensen 


SBCM 


14 June 


1892 


6 


ES Bryant 


WFVZ* 


21 May 


1895 


9 


WF Hill 


FMNH 


12 June 


1895 


12 


RS Judd 


PMYU 


23 May 


1898 


15 


PB Peabody 


WFVZ 


21 May 


1900 


13 


F Maltby 


NYSM 


31 May 


1901 


13 


HKJob 


USNM 


15 May 


1902 


— 


DR Ducks 


UMMZ 


20-23 May 


1902 


— 


DR Ducks 


UMMZ 


18 May 


1902 


16 


NA Francis 


MCZ 


23 May 


1902 


13 


NA Francis 


MCZ 


29 May 


1904 


9 


AD Doerge 


SDNHM 


26 May 


1909 


14 


RS Judd 


PMYU 


15 May 


1916 


15 


EJ Booth 


MVZ 


18 May 


1917 


16 


GC Withey 


WFVZ 


18 June 


1917 


13 


GA Withey 


DMW 


22 May 


1923 


14 


GC Withey 


UWGB 


24 May 


1923 


12 


GC Withey 


WFVZ 


3 June 


1923 


10 


DOgg 


CM 


26 May 


1924 


13 


GF Abbey 


WFVZ 


28 May 


1924 


9 


GC Withey 


WFVZ 


28 May 


1924 


7 


GC Withey 


SBCM 


28 May 


1924 


16 


GC Withey 


WFVZ 


31 May 


1924 


10 


GC Withey 


UWGB 


31 May 


1924 


12 


GC Withey 


UWGB 


5 June 


1924 


12 


GF Abbey 


SBCM 


27 May 


1925 


9 


GL Davy 


PMYU 


31 May 


1925 


14 


GA Withey 


MVZ 


19 May 


1926 


14 


GL Davy 


WFVZ 


21 May 


1926 


10 


GA Withey 


WFVZ 


11 May 


1927 


13 


GA Withey 


DMW 


1 2 June 


1927 


11 


GL Davy 


WFVZ 


12 May 


1928 


9 


GA Withey 


WFVZ 


18 May 


1928 


15 


GA Withey 


WFVZ 


1 June 


1929 


12 


GA Withey 


SBMNH 


19 May 


1930 


14 


GC Withey 


WFVZ 


25 May 


1930 


14 


GC Withey 


PMYU 


6 June 


1930 


11 


GC Withey 


PMYU 


6 June 


1930 


11 


GC Withey 


PMYU 


20 May 


1931 


10 


GC Withey 


USNM 


24 May 


1931 


12 


GC Withey 


WFVZ 


24 May 


1931 


13 


GC Withey 


NYSM 


31 May 


1931 


12 


GL Davy 


WFVZ 


5 June 


1931 


8 


GC Withey 


WFVZ 


14 May 


1932 


13 


GC Withey 


WFVZ 


20 May 


1932 


12 


GC Withey 


WFVZ 


12 June 


1932 


13 


GC Withey 


SBMNH 


18 May 


1933 


9 


GC Withey 


CU 


26 May 


1933 


11 


GC Withey 


CMNH 


13 May 


1934 


12 


GC Withey 


WFVZ 


8 June 


1934 


10 


GC Withey 


WFVZ 


I'll 


1934 


11 


GC Withey 


Minot 






285 






777 


1871 


8 


FV Hayden 


USNM 


17 May 


1879 


9 


GS Agersborg 


MCZ 


26 May 


1882 


8 


HB Bailey 


AMNH 


27 May 


1882 


8ne 


HB Bailey 


AMNH 
Continued 



I 



2002 



Houston: Greater Prairie-Chicken on the Canadian Prairies 



Table 1. Continued 



# sets sets 


eggs/set 










ceggs 












Locality 


Day/Mo 


Year 


#eggs 


Collector 


Institution 


Vermillion 


30 May 


1882 


8 


HB Bailey 


AMNH 


Vermillion 


30 May 


1882 


7 


HB Bailey 


AMNH 


Flandreau, Moody Co. 


2 June 


1882 


11 


HB Bailey 


WFVZ 


Egan, Moody Co. 


16 June 


1882 


7 


(via BF Goss) 


MPM 


Vermillion 


16 June 


1882 


8 


HB Bailey 


AMNH 


Vermillion 


23 June 


1884 


10 


GS Agersborg 


CAS 


Roswell, Miner Co. 


May 


1888 


12 


FA Patton 


UMA 


Sanborn Co. 


7 May 


1889 


10 


FA Patton 


WFVZ 


Miner Co. 


May 


1889 


11 


FA Patton 


AMNH 


Roswell, Miner Co. 


14 May 


1889 


12 


FA Patton 


HSU 


Miner Co 


17 May 


1890 


12 


A Hewitt 


WFVZ 


Harrison 


18 May 


1890 


10 


WC Colt 


MVZ 


Miner Co. 


10 May 


1891 


14 


FA Patton 


USNM 


Harrison 


25 May 


1891 


12 


WC Colt 


MVZ 


Roswell 


2 June 


1891 


11 


BV Jones 


BMNH 


Bryant 


20 May 


1895 


15 


HE Lee 


USNM 


Watertown 


25 May 


1895 


10 


GW Dixon 


WFVZ 


Sanborn Co. 


14 May 


1898 


15 


FA Patton 


WFVZ 


Hamlin Co. 


13 May 


1903 


15 


HE Lee 


OMNH 


Hamlin Co. 


16 May 


1903 


14 


HE Lee 


MVZ 


RondelL Brown Co. 


18 May 


1905 


10 


J Collins 


WFVZ 


Armour, Douglas Co. 


6 May 


1911 


10 


A Walker 


WFVZ 


Sanborn Co. 


20 May 


1914 


— 


FA Patton 


Earlham 


Sanborn Co. 


1 June 


1922 


15 


FA Patton 


UAM 


Grand Total 131 119 


11.10 




1321 







Notes: In 7th vertical column. ^^ indicates that a single egg was obtained from an oviduct. Egg numbers in parentheses, 
from incomplete sets, are not counted in totals. In 8th vertical column, ne = eggs no longer extant; otherwise, ne means 
northeast. In 10th vertical column, WFVZ* indicates set came from Cumberland Science Museum, Nashville Tennessee. In 
some clutches in some collections, some of original eggs are no longer extant, but are still counted: e.g.. Stony Plain, 
Alberta, there are now only 1 1 eggs remaining from original set of 14. Institution abbreviations in 8th vertical column can 
be found alphabetically in Acknowledgements. 



H. H. Mitchell reported it as "fairly common resident 
in transition zone, apparently extending its range 
northwestward," and Bradshaw's annual report of 
the game commissioner said it was now "reported 
from almost all points where the [Sharptail] occurs." 
Andrew Holmes reported "several fairly large cov- 
eys" at Davis and Henribourg, south of Prince 
Albert, in the fall of 1924 (Bradshaw 1925). 

Spread into Alberta (Figure 3) 

No reliable dates exist for the arrival of the 
Greater Prairie-Chicken in Alberta; it was never 
common there. Whether the "old-timers" correctly 
remembered it in the late 1890s, as told to Rowan, 
seems highly doubtful (Mitchell 1959). However, by 
1911 and 1912 it was "common" in the Hanna dis- 
trict (Mitchell 1959), and several were shot in east- 
ern Alberta in the fall of 1913 (Lawton 1913). Thus 
in 30 years it had spread west 1000 km from 
Winnipeg, at an average rate of about 33 km per 
year, doubling the speed of its Minnesota - North 
Dakota spread. The first specimen from the Red 
Deer area was shot near Buffalo Lake, east of 



Mirror, by George Cook on 26 December 1914 
(Horsbrugh 1918). Single individuals were shot near 
Bawlf in 1916 (Farley 1932), in the Delia area north- 
east of Drumheller in 1918 (Mitchell 1959), and 
south and east of Camrose in October 1 926 (Farley 
1932). It was hunted near Calgary and Edmonton, 
and seen north to Lac la Biche (Rowan 1926; Salt 
and Salt 1976). It nested near Buffalo Lake. Three 
nests were found at the south end of Beaverhill Lake, 
17-18 May 1924 (Salt and Wilk 1958; Farley 1932; 
Mitchell 1959; Lister 1979). Three or perhaps four 
booming grounds near Beaverhill Lake were in use 
in 1925. William Rowan's field notes show it was 
also common at Gough Lake and east of Big Valley, 
1920-1925. In a letter to George J. Mitchelldated 15 
April 1957. Rowan told of "breeding populations"at 
Sullivan Lake in 1924, and also at Wavy Lake east 
of Daysland. Salt and Wilk (1958) mapped only nine 
known localities of occurrence in Alberta. 

Stable period in Saskatchewan, 19 10-1 920s 

The open, fertile plains, quickly broken and plant- 
ed to wheat and other grains, interspersed with sum- 



f 



10 



The Canadian Field-Naturalist 



Vol. 116 



Table 2. Main Collectors of Greater Prairie-Chicken egg 
sets Northern Great Plains 



Collector 


#sets 


State, Prov 


Years 




G. C. Withey 


24 


ND 


1917 


1934 


C. P. Forge 


16 


MB 


1900 


1905 


G. A. Withey 


7 


ND, SK 


1927 


1929 


G. L. Davy 


6 


ND, SK 


1925 


1927 


F. A. Patton 


7 


SD 


1888 


1922 


H. B. Bailey 


6 


SD 


1882 




G. F. Abbey 


4 


ND, SK 


1924 


1925 


H. H. Pittman 


3 


SK 


1923 


1925 



First egg set. 1879, Vermilion, South Dakota. G. S. 
Agersborg. Last set, 1934, McHenry Co., South Dakota, 
G. C. Withey, 43 sets in Western Foundation of Vertebrate 
Zoology. 50 museums and university collections contacted. 
36 had Northern Great Plains egg sets. 



merfallow every second or third year, soon offered 
insufficient cover for Greater Prairie-Chicken in the 
flattest and most intensively farmed areas. 

Regions that remained attractive were grassy, low- 
lying, somewhat moist areas such as those around 
Old Wives Lake, the north end of Last Mountain 
Lake, and the Quill Lakes, together with areas in the 
Qu'Appelle Valley. H. H. Mitchell sighted them at 
Craven on 15 February 1916 and at Imperial Beach 
on 24 October 1922 and collected specimens at 
Imperial Beach on 27 October 1922 (two), and 21 
October 1924. H. Eutenner (Euteneir?) shot one at 
Holdfast in 1920 and presented it to Mitchell at the 
museum; a specimen from R. A. McEwen at 
Nokomis had no date. (Belcher 1961). They were 
still present southwest of Snipe Lake until 1926, 
when Elizabeth [Hubbard's] family moved to 
Grenfell (Hubbard 1976). Several birds were seen 
near Big Quill Lake at Dafoe on 10 and 11 October 
1930 by Bradshaw and Bard (Belcher 1961). Four 
were seen at the north end of Last Mountain lake by 
F. G. Bard on 5 June 1936 and J. D. Soper saw a sin- 
gle bird at Watrous on 5 July 1936 (Soper 1970). 

In the Weyburn and Virden map sheet areas in 
southeastern Saskatchewan, they were "common and 
widespread" from 1910 to at least 1920. At Percival, 
John Nelson's oldest brother shot one about 1928. 
Ralph Stueck at Abernethy and Ken Skinner at 
Katepwa Lake saw them regularly in the 1920s but 
they disappeared in the late 1930s (Callin 1980). 
R. D. Symons (1967) observed a booming ground 
near Moosomin, presumably in the 1920s. 

In the Moose Jaw area, sportsmen found them 
common through at least the early 1930s. Hugh 
McCrea had sightings at Old Wives Lake in the early 
1930s and Tom Beveridge had them at Mortlach in 
1934, where they persisted until at least early 1942 
(Knight 1967). In the Allan Hills, where he farmed 
in the upper Arm River valley northeast of 



Davidson, 1920-1925, R .D. Symons heard the 
"booming, hollow note ... in those crocus-sweet 
spring mornings ... awakened ... from sleep at day- 
break" (Symons 1967). 

Along the United States boundary a few were still 
present in 1927: one near Altawan, 8 August, several 
at Lonesome Butte, south of Lafleche, 9-17 
September, and two at Buffalo Gap south of 
Willowbunch on 10 September (Soper 1970). 

Montana 

If this species indeed migrated south (or south- 
east?) in winter and if the 1965 and 1972 sightings in 
extreme southern Alberta originated from Montana, 
one would expect records across northern Montana, 
south of Alberta and western Saskatchewan. Such 
records do not exist. Saunders' state list in 1921 
recorded only a single specimen from a most unex- 
pected location, deep within the state and not even in 
good prairie chicken habitat (Bob Eng, personal 
communication), shot by John R. Bane near Huntley, 
northeast of Billings, in the fall of 1917 (Saunders 
1921). P. D. Skaar (1987) doubted the validity of this 
record, believing that Saunders had not seen the 
specimen. If valid, might it have reached Huntley on 
a railroad flat car? 

Only in the extreme northeast comer of Montana 
(Sheridan County) have there been credible records 
at any time. The first record is for the sandhills 40 
km north of Culbertson, west of the present hamlet 
of Medicine Lake, in 1904, when J. B. Lyons shot 
over 200 'squaretails' in four days. A hunter shot 
two south of the hamlet of Homestead, 10 km farther 
south, about 1919. Joseph A. Morin saw them in the 
Wolf Creek area southwest of Plenty wood in the fall 
of 1936. That year, William T. Krummes, assistant 
manager of the Medicine Lake National Wildlife 
Refuge (NWR), listed them as rare, but included a 
photo of a booming male just north of Medicine 
Lake. Frank Reuter, a rancher near Medicine Lake, 
had a large booming ground in a hay meadow until 
about 1940-41. Farther south, in the Yellowstone 
valley near the hamlet of Intake between Glendive 
and Savage, Art Suckow gave a description of seven 
or eight seen in early June 1944, the last credible 
record in Montana (Walcheck 1980). A careful sur- 
vey of Sheridan County by Bob Eng (personal com- 
munication) in 1952 revealed no sight or sound of 
Greater Prairie-Chickens, but after a hiatus of 23 
years there was a single unconfirmed report in 
1961-62. 

North Dakota 

In North Dakota, the arrival and distribution are 
well documented, after it had "followed the plow ... 
five hundred miles ... in fifty years ... an average of 



2002 



Houston: Greater Prairie-Chicken on the Canadian Prairies 



11 



ten miles a year" (Partch 1973). The Greater Prairie- 
Chicken was first reported at Grand Forks in 1880, 
and had a "booming ground" in the most northeaster- 
ly county, Pembina, by 1882. By 1884, it was com- 
mon at Fargo and had moved 100 km west of there, a 
spread of 500 miles (800 km) in 50 years. By 1900, 
this species had spread westward almost exactly to 
the Montana boundary; it was 40 km short of the 
Montana boundary at Medora; 27 km from it at 
WiUiston; 8 km from it at Marmarth and 3 km from 
the boundary at Beach (Walcheck 1980); at that time 
it occupied all but the extreme southwestern comer 
of North Dakota. 

Between 1917 and 1934, enthusiastic oologists, 
David Ogg, his son-in-law, George L. Davy, and his 
friend George A. Withey, collected 37 clutches in 
McHenry and Bottineau counties, especially near 
Antler, North Dakota, which is just a few km south 
of the junction of Manitoba and Saskatchewan with 
the 49th parallel (Table 1, Bechard and Houston 
1984). As late as 1940, roadside counts gave an esti- 
mated North Dakota state population of 430 000 
individuals, falling to 4,000 to 5,000 by 1964 and 
fewer than 400 by 1972 (Stewart 1975). On the 
Arrowwood NWR, 360 Greater Prairie-Chickens and 
2100 Sharp-tailed Grouse were counted in 1941, 
dropping to zero prairie chickens and 250 Sharp- 
tailed Grouse by 1966 (Kirsch et al. 1973). "When 
the proportion of grain to grassland was about 20 to 
80, the Pinnated moved in and began to increase ... 
As the proportion moved up to 50:50, the population 
... increased to tremendous figures ... But then the 
ratio tipped over the other way, and there was more 
land in crop than in native grassland. Then the pin- 
nate could fmd plenty of food but no nesting cover ... 
or escape cover. He was literally plowed out of 
house and home" (Anonymous 1953). By 1997. the 
spring breeding population in North Dakota was 
"about 300 with most living on or near the Sheyenne 
National Grassland" (Kobriger 1999). 

Saskatchewan breeding records 
(including flightless young) 

The first Saskatchewan nest in 1897 has been 
mentioned above. H. C. Grose collected a set of 10 
eggs near Melville on 10 June 1918 (Houston 1993). 
Pittman at Wauchope found nests with eggs in three 
successive years, 1923, 1924 and 1925 (Nero and 
Lein 1971). The enthusiastic oologist group residing 
in Antler, North Dakota, within 3 km of the 
Canadian boundary, collected at least two sets within 
Saskatchewan, and four other sets that were either 
within Saskatchewan or within one mile of it: 1 1 
eggs on 6 June 1925. 9 eggs on 29 May 1927. 10 
eggs on 8 June 1927, and 10 eggs on 14 June 1927 
(Table 1). The sets definitively named as from with- 
in Saskatchewan, 12 eggs from North Portal on 3 
May 1925, and 14 eggs south of Gainsborough on 12 



May 1925, were both collected by George Abbey 
(see Table 1 and also Bechard and Houston 1984). 
At Quill Lake, five chicks were seen 18 June 1929 
by Fred Bradshaw, and a female with a brood was 
seen there by Todd (1947), 20 June 1932. J. P. 
May wood found a nest with 14 eggs, 6 km south- 
west of Nipawin, 22 June 1947. M. G. Street flushed 
a female with seven young near Nipawin on 28 July 
1947 (Houston and Street 1959). Wallace Black had 
several nest records at Somme, 1920-1942 (Hooper 
1992). Peter McLellan found a hen with three young 
in deep grass 3 km east of Areola on 13 September 
1955 (letter, 13 September 1955). The latest report 
was an adult with eight half-grown young in a long 
grass meadow in Cypress Hills Provincial Park, 14 
July 1966 (Pegg 1967). 

Migration? 

Did some Greater Prairie-Chickens migrate? Salt 
and Wilk (1958) explained the absence of winter 
sightings in Alberta (but note the 26 December 1914 
specimen above) with the remarkable but unsubstan- 
tiated claim that Greater Prairie-Chickens "left the 
province during the winter." There is reasonable evi- 
dence (Cooke 1885) that females, especially, migrat- 
ed from southern Minnesota and northern Iowa to 
southern Iowa and northern Missouri. However, 
Hamerstrom and Hamerstrom (1973) found no evi- 
dence of migration in their local Wisconsin popula- 
tion. 

Museum egg data sets 

Because of the paucity of other breeding records 
in the five jurisdictions, my wife Mary joined me for 
a week in the Western Foundation of Vertebrate 
Zoology, transcribing 3660 egg data sets for all 
Northern Great Plains species, including 34 Greater 
Prairie-Chicken sets. We also transcribed data in the 
Saskatchewan Museum of Natural History (SMNH) 
and the University of Saskatchewan Biology 
Museum (USask). Next, I wrote to all museums 
reported by Kiff and Hough (1985) to hold egg sets, 
and collated their data (Table 1 ). 

Museum egg data are the best breeding evidence 
extant for many localities in many years. They also 
provide average numbers of eggs per set: 11.5 in 
Alberta (n = 2 ); 10.9 in Saskatchewan (n = 8 ); 10.0 
in Manitoba (n = 30); 1 1.9 in North Dakota (n = 50); 
and 1 1.0 in South Dakota (n = 26) (Table 1). 

Relative numbers 

Relative abundance has several times been stated 
in terms of the numbers of their nearest related 
species. In 1886. near Winnipeg. Greater Prairie- 
Chickens were about half as common as the Sharp- 
tailed Grouse (Nash, in Thompson 1890). Near 
Saltcoats from 1910 to 1920 they were as common 
as the Sharp-tailed Grouse (Baines 1956). a claim 



12 



The Canadian Field-Naturalist 



Vol. 116 



made also by Ferry (1910) at the Quill Lakes in 
1909. In southern Saskatchewan, they were the 
"commonest upland game bird ... the real open 
prairie dwellers" at some unspecified time after 1905 
(Taylor 1955). Rutherford (1914) claimed that 
"grouse, pinnated, ruffed and sharptail, are to be had 
in immense numbers." In adjacent North Dakota, 
E. T. Judd (1927) found that the Greater Prairie- 
Chicken was "practically unknown" in the early 
1890s, but increased after 1910 to become common 
by 1915: "if it continues to increase as it has in the 
past five years it can soon be classed as an abundant 
bird," to a large extent replacing the Sharp-tailed 
Grouse. Near Raymore, Saskatchewan, it increased 
rapidly in three years to become equivalent in num- 
bers to the Sharp-tailed Grouse about 1920 (Charles 
Harris in W. C. Harris, unpublished manuscript). 

Ten-year population cycle 

Greater Prairie-Chicken did not persist long 
enough on the Canadian prairies for a population 
cycle to become evident. Elsewhere, Minnesota pop- 
ulations peaked in 1863, 1871, 1880-81, and 
1894-95, and a century later in 1982 and 1992; in 
Wisconsin they peaked in 1940, 1950, and 1970-71 
(Svedarsky et al. 1997). This approximate ten-year 
cycle is similar to that of the Great Horned Owl, 
Snowshoe Hare, Lynx, and to that of other upland 
game birds, the Ruffed Grouse, Sharp-tailed Grouse 
and Spruce Grouse elsewhere in North America 
(Houston et al. 2002, in press). 

Habitat requirements 

"...wildlife abundance or scarcity is fundamentally 
a function of habitat suitability. ... nest-brood cover 
[is] the weak link in the Prairie Chicken life chain" 
(Olson in preface to Hamerstrom and Hamerstrom 
1973). Roberts (1936) postulated that the presence of 
grain for food was the stimulus for the westward and 
northwestward spread within the United States; the 
Greater Prairie-Chicken entered new areas soon after 
grain was planted. The importance of grains was cor- 
roborated by a 1962 study in Missouri, where 44% 
of diet volume was composed of com, soybeans and 
sorghum, in descending order of importance, with 
wheat and oats together contributing another 9% 
(Korschgen 1962). 

Macoun (1900) described it as a "true prairie bird 
as observers speak of it always being found in the 
open even in the severest weather." Everywhere it 
tended to choose low-lying areas with high grass. 
Schroeder and Robb (1993) point out that its pre- 
ferred habitats tend to be interspersed with or sur- 
rounded by oak trees, reminiscent of the Passenger 
Pigeon (Ectopistes migratohus). 

As it followed the plow westward, the Greater 
Prairie-Chicken presumably had the best of both 
worlds, but only for a short while. Long grass 



offered cover and grain fields provided food, the two 
habitats often "interdigitating" (Johnsgard and Wood 
1968). They seemed to take advantage of the thick 
grass that grew luxuriantly between the time of less- 
ened grazing pressure due to the demise of the Bison 
and the subsequent secondary over-hunting of 
Pronghorn (Antilocapra americana), and the later 
increase in cattle (graphed by Johnston and Smoliak 
1976). The grassland range was "not fully stocked 
until about 1920." Suppression of grass fires by set- 
tlers was a further benefit. 

Next, as even more land was cultivated, especially 
when cultivation reached 60% of the land area, and 
as ever-fewer parcels of grass interspersed with rose- 
hips (Judd 1905) had sufficient area to support a 
group of prairie-chickens, this species left (Rue and 
Allen 1973). And, in the absence of fire, invasion of 
grasslands by trees and shrubs made grassland frag- 
ments even smaller. In North Dakota, the ideal habi- 
tat was grass 20 inches (50 cm) in height and suffi- 
ciently dense to completely conceal a nesting 
female; fields annually grazed, annually hayed, or 
idle >10yr were undesirable (Kirsch 1973). In 
Oklahoma, Jones (1963) noted that mean height of 
vegetation in booming grounds was 15.1 cm., at nest 
sites 45 cm (range 25 to 70 cm), and escape cover, 
60 cm. 

The drought of the 1930s, complicated by over- 
grazing, sounded the final knell; few coveys could 
find luxuriant grass to provide adequate cover on the 
Canadian prairies. Between 1920 and 1940 there was 
also a slowly increasing shift from harvesting with 
threshing machines (producing attractive straw- 
stacks) to combines (Svedarsky et al. 1997); the 
elimination of strawstacks was not complete until 
1960, after the demise of the Greater Prairie- 
Chicken. 

Decline in Alberta 

In Alberta, Greater Prairie-Chickens had "practi- 
cally disappeared" by the mid- 1930s. The last sight- 
ing in the Drumheller area was by W.R. Salt north of 
Hand Hills, east of Drumheller, in 1925 (Kondla et 
al. 1973). They disappeared from the lek at the south 
end of Beaverhill Lake about 1929 (Farley 1932), 
except for a single in May 1932 and three seen there 
24 May 1934 (Lister 1979). 

Decline in Saskatchewan 

In Saskatchewan, during the First World War all 
grouse numbers, including those of the Sharp-tailed 
Grouse and Ruffed Grouse, collapsed to such a 
degree that the season was closed for Sharp-tailed 
Grouse and Greater Prairie-Chicken from 1916 
through 1920. Neil Gilmour, Game Guardian at 
Moose Jaw, reported in 1923 that the Sharp-tail and 
the Ruffed Grouse had rebounded and continued to 
increase, but the Pinnated, once fairly common, was 



2002 



Houston: Greater Prairie-Chicken on the Canadian Prairies 



13 



I 



not coming back in numbers as the other two species 
had done (Bradshaw 1923). 

The decline in Greater Prairie-Chicken numbers 
began early. They were common north of B lad worth 
until about 1918, with two booming hills of 12-15 
birds each (Roy 1996). They were last observed 
booming on knolls near Langbank in 1924-1927 by 
Geoffrey Hewson (Belcher 1961; Hewson 1977), 
and last seen near Sheho about 1926 by William 
Niven (Houston 1949). They disappeared from the 
Storthoaks area by the late 1920s, certainly by 1935 
(Stelfox 1980; Ray Barber, personal communica- 
tion). Last "booming ground" at Cut Knife was in 
1931, and the last in a hunter's bag at Maidstone was 
in 1932 (Symons 1967). Near Davidson and 
Imperial, Saskatchewan, they were "greatly 
decreased in numbers" by 1932, although there were 
still three booming grounds near Davidson and 
Stalwart and east of Imperial; two were seen at 
"Devil's lake" west of Simpson, and five specimens 
were collected at Last Mountain Lake (Todd 1947). 
They disappeared from the Qu'Appelle valley in the 
late 1930s (Callin 1980). The last observation at 
Raymore was of a single male observed by A. 
Cameron dancing with Sharp-tailed Grouse in the 
springs of 1947 and 1948 (Wayne C. Harris, unpub- 
lished manuscript). 

There are only four known observations within 
the Saskatoon area. A mounted specimen in the pos- 
session of N. Loucks, Main Street, Saskatoon, had 
been collected about 1 February 1925 (interview 
with J. B. Gollop about 1970). A male specimen in 
the University of Saskatchewan Biology Museum, 
was collected at or near Saskatoon, 12 October 1931 
(Smith 1968). From 1935 to 1945, Judge A. H. 
Bence (1945) observed the "odd covey" north of 
Saskatoon and he once watched the courtship 
"booming" display of the male near a slough 
between Borden and Radisson. In 1946, while col- 
lecting specimens for the Royal Ontario Museum, 
Farley Mowat (1946) flushed one near Dundurn, on 
the road to Proctor Lake, on 21 May [date obtained 
from Mowat' s field notes]. 

We can only guess how common Greater Prairie- 
Chickens were in extreme southern Saskatchewan, 
but a few persisted until 1940 and 1941. Soper saw 
one or two every day at Rock Creek, south of 
Lafleche, on 7 and 8 June; one near Wood Mountain 
on 9 June 1940, and one along the Frenchman River 
below Val Marie on 1 6 June and one farther west at 
Middle Creek on 20 June 1941 (Soper 1970). 

Probably Greater Prairie-Chickens were never 
common near the northern edge of their maximal 
range, though paradoxically this was where some of 
the final sightings occurred; here the requisite long 
grass persisted in clearings along the fringe of low- 
lying areas in mixed forest. The last sighting at 
Prince Albert was about 1932 (Brooman /// Houston 



and Street 1959). Maurice Street or his brother 
Stanley saw a flock of nine, 8 km east of Nipawin, 
10 December 1935. They were regular at 
Kelvington, Lintlaw and Somme until 1934, with the 
last sighting at Somme about 1942 (Hooper 1992). 
Several were seen along the west boundary road of 
Prince Albert National Park on 14 July 1940 in an 
area with small patches of prairie or grassy wood- 
land glades ... a surprise (Soper 1952). James 
McCunn, on his farm a mile east of Codette, saw and 
shot his first on 20 September 1945. Two months 
later, on 20 November 1945, McCunn saw a flock 
feeding at a granary 43 km east of Nipawin (Houston 
and Street 1959). At Emma Lake, a flock of 12 in 
natural or "moose" meadows in 1937-38 decreased 
to three in 1939, the year that Mowat saw single 
individuals in Prince Albert National Park and at 
Montreal Lake (Mowat 1947). The last record in 
central Saskatchewan was of a lone bird 5 km north 
of Snowden, 10 June 1950 by Walter and Billy 
Matthews and three other observers (Houston and 
Street 1959). 

In southern Saskatchewan, Soper' s last sightings 
were one near Reed Lake, just south of Morse, on 16 
July 1946. a pair near Redvers on 8 August 1946; 
two south of Bender post office, south of 
Whitewood, on 10 August 1946; and one near Red 
Jacket on 20 September 1947 (Soper 1970). 

Decline in Manitoba 

Having peaked in numbers, the decline was evi- 
dent early at Birtle, Manitoba, where it was only 
"abundant until 1916" (Bird 1930). Bruce Noton 
(personal communication) remembers seeing two in 
the ditch en route to Royal School, 6 km west and 
2 km north of Boissevain, about 1931; his friend, 
Herb Patterson, shot two south of Wawanesa in the 
early 1930s. In the winters of 1933 and 1934, Victor 
Latta reported a single bird near Shelley. T. 
Schindler shot his last one near Clandeboye about 
1934 (Gardner 1981). G. W. Malaher saw over 300 
within 100 km of Winnipeg as late as 1936 
(Hamerstrom 1956). Douglas Shanks told of small 
numbers and at least one booming-ground near Old 
Pinawa in the late 1930s; his last sighting was of 
four in the same area in the winter of 1941-42 
(Taylor 1983). The Greater Prairie-Chicken disap- 
peared from Lyleton in the 1940s (Knapton 1979) 
and from the Oak Hammock Bog in 1946 (P. 
Romanic in Gardner 1981), but was present along 
the south edge of Delta Marsh "until 1950 when it 
declined rapidly." (Hochbaum 1971). Three were 
seen at Windfield Swamp near Stonewall by Robert 
E. Jones in the winter of 1950-51 (personal commu- 
nication). John H. More saw a Pinnated Grouse with 
a group of Sharptails near Russell, on 8 October 
1953 (letter, 19 October 1953). 



14 



The Canadian Field-Naturalist 



Vol. 116 



Causes of decline 

Bird (1961) ascribed the decline to two factors: 
over-shooting and destruction of nesting sites by 
intensified agriculture. A review article reported "It 
is ironic that the Greater Prairie-Chicken at first 
prospered from an expanding agriculture ... [until] 
the further development of cereal farming and cattle 
ranching eliminated the tall grass vegetation on 
which the species had depended.... the late 1870s and 
early 1880s were wetter than normal and grass grew 
luxuriantly" (Johnston and Smoliak 1976). Godfrey 
(1986) summarized the situation similarly: "In the 
interim between the disappearance of the bison from 
the prairies (about 1880) and the beginning of inten- 
sive settlement (about 1920), the natural prairie 
grasslands flourished as never before and probably 
never will again. Natural grassland is ideal for the 
Greater Prairie-Chicken and it too flourished in that 
period and declined rapidly when its habitat was 
destroyed by the settlements of man." 

Christisen (1969) has provided a list of factors 
detrimental to this species, with special reference to 
the United States: (a) conversion of grass to tilled 
crops, (b) removal of grass cover by grazing and 
haying, (c) overgrazing, (d) extensive burning, (e) 
loss of open grasslands to shrubs and trees, (f) 
drought in the 1930s, and (g) lack of winter food. 
Remaining populations in the 1960s were confined 
to areas with some Big Bluestem Grass, and in 
Missouri required areas with 25 to 30% grass. 
Another factor contributing to decline in Illinois was 
the decrease in production of redtop, Agrostis alba, 
which gave excellent nesting and brood cover, but 
was largely replaced by legume hays which were 
often harvested at the time when broods are hatching 
(Yeatter 1963). Depredations by raptors, an ever- 
present threat, seem not to have been a serious factor 
in the decline (Berger et al. 1963). Burning was not 
always detrimental; use of grassland increased the 
second year after a spring bum (Westemeier 1973). 

Robel (1970) reported from Kansas that of 60 
Greater Prairie-Chicken males on three booming 
grounds, 2.7 km apart, only 33 took up territory and 
only six (two at each booming ground) in any one 
year were involved in mating. Whether or not such 
social systems are dysfunctional in declining popula- 
tions seems not to have been studied. 

Skinner (1974) found that the two small colonies 
persisting in northern Missouri in 1973 had suffered 
mainly from conversion of grassland to cropland and 
intensive grazing on what pastures remained. Winter 
and summer, prairie-chickens preferred cover of 8 to 
10 inches in height, hence Skinner suggested as a 
management technique for this species that the 
remaining "warm season grasses" should "never be 
grazed below ten inches"! Ryan et al. (1998) found 
that contiguous prairie > 65 ha supported a stable 
population over 27 years in southwestern Missouri, 



whereas nest success was lower in the prairie mosaic 
landscape and the population there declined. 

Interbreeding/hybridization 

None of the authorities quoted above mentioned 
hybridization as a contributing factor to the decline. 
However, hybridization with the Sharp-tailed Grouse 
(Evans 1966; Rowan 1926; Sparling 1980; Lumsden 
1970), and less commonly with the Ring-necked 
Pheasant (Lincoln 1950), has had deleterious effects. 
On Manitoulin Island, Ontario, colonized by Sharp- ■ 
tailed Grouse from the north, and by Greater Prairie- | 
Chicken from the south (Lumsden 1949), hybridiza- 
tion seems to have led to the complete disappearance 
of the Greater Prairie-Chicken (Lumsden 1970; 
Godfrey 1986). 

Grouse interbreed remarkably freely. Alberta's 
first two hybrids of Greater Prairie-Chicken with 
Sharp-tailed Grouse were reported from Gough Lake 
in 1918 and from near Edmonton in 1925 (Rowan 
1926). C. G. Harrold (1933) identified two hybrids 
with a party of Sharptails near Old Wives Lake in 
May 1922, but Saskatchewan's first recorded hybrid 
specimen was not taken until 22 October 1933 by 
J. C. Lusted south of Regina at Truax (Belcher 
1961). Some, perhaps most, of the last records of 
Greater Prairie-Chickens in the prairie provinces 
were in company with Sharp-tailed Grouse or Sage 
Grouse. At Raymore, Wayne C. Harris (personal 
communication) saw a hybrid Sharp-tailed Grouse 
X Greater Prairie-Chicken on 25 June and again on 
24 December 1971. On 20 April 1987, Wayne C. 
Harris and Don Weidl saw a female Greater Prairie- 
Chicken on a Sage Grouse dancing ground between 
Killdeer and Val Marie in the west block of the 
nascent Grasslands National Park (Gollop 1987, 
Harris et al. 1987). Three days earlier they had seen ■ 
what was hypothesized to be the first-ever hybrid 
Sage Grouse X Greater Prairie-Chicken. However, 
Dale Hjertaas (1995) photographed a Sage Grouse 
X Sharp-tailed Grouse hybrid south and west of 
Lafleche in May 1988, a hybrid similar in appear- 
ance to that reported earlier by Harris farther south. 
Similar hybrids have been reported from Montana 
(Eng 1973) and Alberta, with DNA analyses as proof 
for the latter (Aldridge et al. 2001). No Sage Grouse 
X Greater Prairie-Chicken hybrid specimen has yet 
been collected, probably due to the minimal overlap 
in their distribution. 

In Nebraska, extensive surveys in the 1960s 
showed that up to 17% of display grounds were 
mixed, and that the minimum rate of Greater Prairie- 
Chicken hybridization varied from 0.3 to 1.2% 
(Johnsgard and Wood 1968). In extreme northwest- 
ern Minnesota, 3 of 1 1 stable display grounds were 
mixed, and up to 3.5% of the birds were hybrids 
(Sparling 1980). 



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Houston: Greater Prairie-Chicken on the Canadian Prairies 



15 



We have no evidence incriminating the introduced 
Ring-necked Pheasant {Phasianus colchicus) with 
the dechne of the Greater Prairie-Chicken in Canada. 
Widespread release of pheasants from the Beaver 
Creek Pheasant Farm south of Saskatoon, 
Saskatchewan, began only in 1946, after the demise 
of the prairie chicken. In Illinois, the relationship is 
more compelling: "the closure of hunting seasons on 
Greater Prairie-Chickens coincided with the initia- 
tion and expansion of pheasant stocking programs. 
...remnant flocks exist outside the contiguous range 
of the pheasant." Further, parasitism of prairie chick- 
en nests by hen pheasants was observed each year 
and parasitized nests were less successful than non- 
parasitized nests. With mixed clutches the pheasants 
hatched a day earlier, and the prairie chicken hen 
would leave "with the parasitic brood before her own 
eggs hatched." A single cock pheasant would domi- 
nate the entire booming ground. Removal of the 
offending cock merely allowed a subordinate succes- 
sor pheasant to assume the same territory (Vance and 
Westemeier 1979). 

In-breeding of isolated remnants on the Canadian 
prairies may have administered the final coup de 
grace. In Illinois, Greater Prairie-Chicken numbers 
declined from several million birds in the mid- 19th 
century (600 000 were marketed in Chicago in 1873; 
Westemeier and Edwards 1987), to a small remnant 
of 46 survivors by 1994. The dramatic crash in num- 
bers is depicted on four maps; extremely low fertility 
and egg success of these isolated survivors was 
attributed to isolation and inbreeding, substantiated 
by a quick recovery of fertility and hatching success 
after 27 1 individuals had been introduced from large 
populations in three other states (Westemeier et al. 
1998). 

Last Alberta specimens 

Shot 1931 between Czar and Hughenden (Mitchell 

1959). 
Shot 1938 near Youngstown (Salt and Wilk 1958). 

Last Alberta sightings 

Mid- 1930s, near the Oldman River west of Fort 

Macleod (Mitchell 1959). 
28 August 1939, two just outside Waterton Lakes 

National Park, not far from the Montana border 

(C. H. D. Clarke in Rand 1948). 

1939, early 1950s, and finally in either 1961 or 1962, 
near Sullivan Lake (Mitchell 1959, Moyles 1987). 

1940, north of Medicine Hat (Mitchell 1959; 
Godfrey 1966, 1986). 

Through 1943, but not in 1944, Dick Beddoes had 
them in his unpublished Christmas bird counts at 
Daysland, southeast of Camrose (Beddoes 1945). 

On 14 September 1965, B.R. Shantz (1966) saw 

I eight birds, 13 km west of Coutts, Alberta, and 



Sadler and Myres (1976) categorized this as 
"questionable." 
5 March 1972, a single bird near Mountain View, 
southwest of Cardston (Salt and Salt 1976). 

Last Saskatchewan specimens 

Shot 30 April 1931 at Imperial Beach by F.G Bard 

(Belcher 1961); 
Shot October 1931 by Dr. J.R. Hoag at Viewfield, 

northwest of Estevan (Belcher 1961); 
Shot 24 Oct 1940 east of Simpson by F.G. Bard 

(Belcher 1961); 
Shot fall of 1942, west of Saltcoats, by Robert Cock 

(Houston 1949); 
Shot Oct 1943 at Marienthal, southwest of Estevan, 

by F.N. Dunk (Belcher 1961); 
Shot 31 Oct 1945 near Carlyle, by R.J. Fyfe 

(Lahrman 1957); 
Shot 3 Nov 1972 near Leader (Hatch 1973). 

Last Manitoba specimens 

Shot in 1957 at Delta Marsh, in the largest remnant 
of true tall-grass prairie, by a group of American 
hunters (Lawrence St. Goddard, personal commu- 
nication, //<ie Robert E. Jones). 

Shot in the fall of 1961 near Langruth (Salt and Salt 
1976; Godfrey 1966). 

Last Manitoba sightings 

1972, a possible sighting in the Broomhill Wildlife 
Management area in southwest Manitoba (Vere 
Hunt Scoii, fide Knapton 1979). 

3 October 1978, Oak Hammock Marsh, Mr. 
McKillop (Gardner 1981). 

1983, Spruce Woods east of Brandon, possible sight 
record by K. Leavesly (Minish 1987b). 

25 May 1986, four near Pilot Mound, observed by 
Doug and Burton Collins (possibly escapes from 
releases made at Grand Forks, North Dakota, near 
the same time; Robert E. Jones, personal commu- 
nication) 

Last Saskatchewan sight records 

Five unpublished Saskatchewan records resulted 
from my 1956 appeal, including the report of Rayner 
(and More's from extreme western Manitoba), 
above. Melvin Adair (letter, 12 March 1956), east of 
Outram, 18 km west of Estevan, believed that Ring- 
necked Pheasants, Sharp-tailed Grouse and Greater 
Prairie-Chicken all suffered severely in the blizzard 
of 1948. When in the spring of 1949 he dug out his 
coyote traps from beneath the melting drifts that had 
been 7.6 m deep, he found the corpses of a covey of 
Pinnated Grouse that had perished in that snowdrift. 
His Ring-necked Pheasant count locally dropped 
from 169 to 1, presumably the result of the same 
storm, reputed to have been the worst since 1888. 



16 



The Canadian Field-Naturalist 



Vol. 116 



Peter McLellan of Areola (letter, 24 March 1956) 
hunted Greater Prairie-Chickens in the Warmley 
area, 18 km north of Kisbey, until about 1936, about 
equal in numbers with Sharptails, and then saw a sin- 
gle individual there about 1946. His brother, J. S. 
McLellan, found them regularly in section 31-8-4w2, 
6 km north and 3 km west of Areola, but a careful 
search revealed none there in 1955, the year that 
Peter found three young closer to Areola. 

Four southern Saskatchewan sightings were near 
lakes. Near Lucky Lake, the last sightings of individ- 
ual birds were on 15 August 1943 and 16 August 
1947 (Roy 1996). Five were seen at Good Spirit 
Lake, 17 January 1946 by J. A. Gunn ("jottings" 
given to the late Mrs. Isabel Priestly, now in posses- 
sion of CSH). Three birds were seen on the Otthon 
flats, southwest of Yorkton, early November 1946, 
by W. D. Lightbody (Houston 1949). J. E. Glennie 
found them northwest of Govan, within a mile of the 
east shore of Last Mountain Lake, about 1950 and 
1951 (letter, 21 May 1956). 

Farther south, David MacDonald of Manor saw 
one in the spring of 1954, three in 1958 and one at a 
Sharp-tailed Grouse dancing ground in the spring of 
1959 (Belcher 1961). Fred Sharp saw three with 
eight Sharp-tailed Grouse, 6 km north of Sidewood 
in September 1959 (Belcher 1961). Roger Tory 
Peterson and James Fisher, arguably the two best- 
known ornithologists attending the American 
Ornithologists' Union meeting in Regina in 1959, 
saw a single cock Greater Prairie-Chicken within a 
mile of the east edge of Old Wives Lake on 25 
August 1959 (Peterson 1960). Two were observed 
carefully near a highway near Stoughton in the fall 
of 1960 (Pringle 1961). Five were carefully 
described near the east edge of Old Wives Lake on 3 
September 1961 (Pratt 1967). One was seen by J. 
Luthi, 5 km north and 13 km west of Simpson on 12 
May 1962 (Luthi 1963). Twelve appeared on the 
farm of Gilbert Johnson near Marchwell on 16 
February 1967 (Johnson 1967). A single individual 
was with six Sharptails near Ruthilda, mid- July 1971 
(Wapple 1977). A hybrid was seen by Wayne C. 
Harris near Raymore, 25 June and 24 December 
1971 (Houston 1971, 1972). Single birds were seen 
near Mortlach on 19 December 1971 and near Caron 
on 16 April 1972 (Brazier 1972a, 1972b). David W. 
Robinson saw two with Sharptails near Avonlea, 20 
September 1975 (Serr 1976). Derek Kreuger saw 
two in the ungrazed section of the northeast quarter 
of Saskatchewan Landing Provincial Park in 1977 
and again in 1978 (Blood and Anweiler 1979). 

Hjertaas et al. (1993) collected additional obser- 
vations for the RENEW report as follows: R. K. 
Brace, a biologist with Canadian Wildlife Service, 
saw one southwest of Lafleche about the summer of 
1966. B. Campbell saw one on a Sharp-tailed Grouse 
lek 18 km south and 13 km east of Mortlach in April 



1968. Don Jackson saw one 12 km north and 4 km 
east of Plunkett in October-November 1972. J. B. 
O'Neil saw one on a roadside 5 km east of Viscount 
in February 1973 and then saw a small flock on three 
consecutive Fridays 7 km east of Plunkett in 
February 1975. Gerhard Stuewe saw a group of 
between six and eight at Teo Lake, west of 
Kindersley, in mid-September 1975. Ross Campbell, 
a hunter, saw one and then three others within 8 km 
of the South Saskatchewan River, south of 
Riverhurst in Game Management Zone 26C in the 
fall of 1983, and then two near Floral in the winter of 
1984-85 through 22 Feb 1985 (Hjertaas et al. 1993). 

Wayne C. Harris saw a female Greater Prairie- 
Chicken in the west block of the National Grassland 
Park (34-2- 12w3) on 20 April 1987 (already men- 
tioned under "Interbreeding/hybridization, above). 
Then on 8 May 1987, Harris saw three Greater 
Prairie-Chicken males on the United States boundary 
just west of the west margin of the east block of the 
park (5-l-7w3); they flushed from an upland area 
and flew into Valley County, Montana, a region 
lacking either historic or recent records of Greater 
Prairie-Chicken. This was the third last report from 
Saskatchewan. 

The second last sighting of probable Greater 
Prairie-Chickens in Saskatchewan were convincingly 
described north of Big Muddy by Carol Bjorklund, 
28 July 1991 (Koes and Taylor 1991). Near the soli- 
tary male were three Sharp-tailed Grouse hens with 
broods (Carol Bjorklund, personal communication). 
Doris Silcox saw a single individual, together with 
25 Sharptails and up to 9 Gray Partridge {Perdix 
perdix) at Carlyle, mid December 1991 through 18 
Feb 1992 (Koes and Taylor 1992); this was the final 
sighting recorded for Saskatchewan and the last for 
Canada. One cannot but wonder how the individuals 
sighted above escaped detection in so many locali- 
ties for so long without being seen, unless they fly 
long distances from the nearest stable groupings in 
the Dakotas. Should that be the explanation, then 
more sightings may occur in future, though without 
appreciable hope of re-establishment anywhere in 
the Canadian west. Indeed, the Greater Prairie- 
Chicken recovery team was disbanded in 1994. The 
species had been listed in Canada as Endangered in 
1978, and then as officially Extirpated in 1990 
(Hjertaas 1998). 

Maps of range 

The original zone of overlap between the Greater 
Prairie-Chicken and the Sharp-tailed Grouse was ini- 
tially narrow (Figure 3 in Johnsgard and Wood 
1968), but as the Greater Prairie-Chicken spread 
westward this zone of overlap greatly enlarged. By 
January 1955, Aldrich and Duvall (1955) showed 
habitat still occupied by Greater Prairie-Chicken in 
southern Manitoba, but none anywhere in 



2002 



Houston: Greater Prairie-Chicken on the Canadian Prairies 



17 



Saskatchewan, and only a small remnant area left in 
central Alberta. By 1979, Westemeier's map showed 
substantial populations persisting only in South 
Dakota, Nebraska, Kansas and Oklahoma. 

Acknowledgments 

I wish to thank Harry Lumsden for unpublished 
infonnation on this species in northwestern Ontario, 
and Ken J. Lungle for copying the 27-page publica- 
tion (1959) by George J. Mitchell, then Game 
Biologist for the Alberta Department of Lands and 
Forests. Wayne Pepper reviewed the annual reports 
of the Saskatchewan Game Commissioner, 
1905-1930. Carol Bjorklund offered data from her 
1991 field notebook. Dale Hjertaas contacted Ray 
Barber of Storthoaks for more detailed information 
beyond that mentioned in Stelfox's 1980 article. 
James R. Jowsey provided information about the 
Saskatchewan egg collection of Ralph and Norman 
Jowsey, now in the Provincial Museum of Alberta, 
Edmonton (PMA). Jocelyn Hudon of PMA provided 
information concerning the John Kowal egg collec- 
tion in their museum. Robert E. Jones, retired upland 
game specialist, provided detailed information con- 
cerning Manitoba sightings. Robert W. Nero provid- 
ed the information compiled from early newspapers 
in southern Manitoba, 1884-1949, prepared by Anita 
Steeg Kovacs. Lany Igl, Dale Hjertaas, Robert W. 
Nero, Allan R. Smith, and Wayne Pepper offered 
constructive criticism. Dan Svedarsky provided three 
recent publications dealing with Greater Prairie- 
Chicken in the northern states. Lloyd W. Kiff pro- 
vided unlimited assistance during our visit to the 
Western Foundation of Vertebrate Zoology (WFVZ) 
and Frieda Kinoshita kindly supplied data on sets 
added to the WFVZ collection subsequently. Marc 
Bechard recorded data sets in the National Museum 
of Canada. The entire project of checking egg collec- 
tions would not have been possible without the 
Inventory of Bird Egg Collections of North America, 
1985, compiled by Lloyd F. Kiff and Daniel J. 
Hough. 

The following 34 curators responded to mail and 
e-mail requests, supplying data re egg sets: 

Tim Armstrong, Adams State College, Alamosa, 
CO (ASC); Emanuel Levine, American Museum of 
Natural History, New York (AMNH): Ann Kessen, 
Bell Museum of Natural History, Minneapolis 
(BMNH); Karen Cebra, California Academy of 
Sciences, San Francisco (CAS); Will Post, Charleston 
Museum, Charleston, S.C. (CM); Starla Miller, 
Clemson University (CU); Tim Matson, Cleveland 
Museum of Natural History, Cleveland, OH (CMNH); 
Gene K. Hess, Delaware Museum of Natural History 
(DMNH); Kirstie M. Bay, Denver Museum of Natural 
History (DM); John Iverson, Joseph Moore Museum 
of Natural History, Earlham College, Richmond, 
Indiana (Earlham); Tom Webber, Florida Museum of 



Namral History, Gainesville (FMG); David Willard, 
Field Museum of Natural History, Chicago (FMNH); 
Tamar Danufsky, Wildlife Department Museum, 
Humboldt State University, Areata, CaUfomia (HSU); 
Nathan Kraucanus, Milwaukee Public Museum 
(MPM); Allen J. Kihm and Rand Rodewald, Minot 
State University (Minot); Alison Pirie and Raymond 
A. Paynter, Jr., Museum of Comparative Zoology, 
Harvard University (MCZ); Carla Cicero, Museum of 
Vertebrate Zoology, University of CaUfomia Berkeley 
(MVZ); Michel Gosselin, National Museum of 
Canada, Ottawa (NMC) [now Canadian Museum of 
Nature]; Joseph Bopp, New York State Museum, 
Albany (NYSM); Gary D. Schnell, Oklahoma 
Museum of Natural History, Norman, OK (OMNH); 
John M. Condit, Ohio State University Museum of 
Biological Diversity, Columbus (OSUM); Fred C. 
Sibley, Peabody Museum, Yale University, New 
Haven (PMYU); Jocelyn Hudon and W. Bruce 
McGillivray, Provincial Museum of Alberta, 
Edmonton (PMA); Selma Glasscock, Rob and Bessie 
Wilder Wildlife Foundation, Sinton, Texas 
(RBWWF); Ross James, Royal Ontario Museum, 
Toronto (ROM); the late Bob Kreba, Royal 
Saskatchewan Museum, Regina (RSM); Robert 
McKernan, San Bernardino County Museum 
(SBCM); Krista A. Fahy, Santa Barbara Museum of 
Natural History (SBMNH); Philip Unit, San Diego 
Namral History Museum (SDNHM); Nancy Glover 
McCartney, University of Arkansas Museum (UAM); 
Katherine L. Doyle, University of Massachusetts, 
Amherst (UMA); Janet Hinshaw, University of 
Michigan Museum of Zoology, Ann Arbor (UMMZ); 
Alan Gubanich, University of Nevada, Reno (UNR); 
William J. Maher, University of Saskatchewan 
(USask); Thomas C. Erdman, University of 
Wisconsin, Green Bay (UWGB); Frieda Kinoshita, 
Western Foundation of Vertebrate Zoology (WFVZ), 
Los Angeles, since moved to Camarillo. 

The following 16 institutions had egg sets of this 
species, but none from the Northern Great Plains: 
Clayton White, Brigham Young University, Provo, 
Utah; Arthur R. Clark. Buffalo Museum of Science, 
Buffalo. New York; Mary Hennen. Chicago 
Academy of Sciences, Chicago, Illinois; Jane E. 
Deisler-Seno, Corpus Christi Museum of Science 
and History. Corpus Christi, Texas; Christine 
Adkins, Cowan Vertebrate Museum, University of 
British Columbia, Vancouver; Steven D. Bailey. 
Illinois Natural History Survey, Urbana. Illinois; 
Kerri Leedy, Oakes Museum of Natural History. 
Messiah College, Grantham. Pennsylvania; Mark B. 
Robbins. Natural History Museum. University of 
Kansas, Lawrence, Kansas; Thomas E. Labedz, 
University of Nebraska, Lincoln, Nebraska; Albert 
G. Mehring, State Museum of Pennsylvania, 
Harrisburg. Pennsylvania; Janice Hall. Putnam 
Museum. Davenport. Iowa; David Denton and 



18 



The Canadian Field-Naturalist 



Vol. 116 



Roseanna Humphrey, University of Colorado 
Museum, Boulder, Colorado; Gary Shugart, 
University of Puget Sound, Tacoma, Washington; 
Edward Marks, Museum of Natural History, Univer- 
sity of Wisconsin Stevens Point, Wisconsin. Louis 
Levene, Cumberland Science Museum, Nashville, 
Tennessee; reported that their one North Dakota set 
had been de-accessioned to the Western Foundation 
of Vertebrate Zoology. Diane Fell reported that the 
Oklahoma Zoo no longer has an egg collection. 

Literature Cited 

Aldrich, J. W., and A. J. Duvall. 1955. Distribution of 
American gallinaceous game birds. United States Fish 
and Wildlife Service Circular 34. 30 pages. 

Aldridge, C. L., S. J. Oyler-McCance, and R. M. 
Brigham. Occurrence of Greater Sage-Grouse X Sharp- 
tailed Grouse hybrids in Alberta. Condor 103: 657-660. 

Anonymous. 1953. National wildlife week, March 15-21. 
North Dakota Outdoors 15(9): 8-23. 

Baines, K. E. 1956. The ups and downs of game at 
Crescent Lake. Blue Jay 14: 65-66. 

Bechard, M. J., and C. S. Houston. 1984. North Dakota 
oologists. Blue Jay 42: 176-183. 

Beddoes, R. 1945. Bird notes. Blue Jay 3: 24. 

Belcher, M. 1961. Recent records of the Greater Prairie 
Chicken in Saskatchewan. Blue Jay 19: 76-77. 

Bence, A.H. 1945. [Square-tailed Grouse]. Blue Jay 3: 
39. 

Berger, D. D., F. Hamerstrom, and F. N. Hamerstrom, 
Jr. 1963. The effect of raptors on prairie chickens on 
booming grounds. Journal of Wildlife Management 27: 
778-791. 

Bird, R. D. 1930. Biotic communities of the aspen park- 
land of central Canada. Ecology 11: 356-442. 

Bird, R. D. 1961. Ecology of the aspen parkland of 
Western Canada in relation to land use. Canada Depart- 
ment of Agriculture Pubhcation 1066, Ottawa. 155 pages. 

Blood, D. A., and G. G. Anweiler. 1979. Resource inven- 
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Received 18 May 2000 
Accepted 10 April 2002 



Status of Common Eiders, Somateria mollissima. Nesting in the 
Digges Sound Region, Nunavut 

J. Mark Hipfneri, H. Grant Gilchrist^, Anthony J. Gaston^ and David K. Cairns^ 

'Biopsychology Programme, Memorial University of Newfoundland, St John's, Newfoundland A IB 3X9 Canada 
^Canadian Wildlife Service, Suite 301, 5204 - 50th Avenue, Yellowknife, Northwest Territories XI A 1E2 Canada. 

E-mail: grant.gilchrist@ec.gc.ca [Author for correspondence] 
^Canadian Wildlife Service, 100 Gamelin Boulevard, Hull, Quebec KIA 0H3 Canada 
"^Department of Fisheries and Oceans, Box 1236, Charlottetown, Prince Edward Island CIA 7M8 Canada 

Hipfner, J. Mark, H. Grant Gilchrist, Anthony J. Gaston, and David K. Cairns. 2002. Status of Common Eiders, Somateria 
molissima, nesting in the Digges Sound region, Nunavut. Canadian Field-Naturalist 116(1): 22-25. 

There is little information available on status of Common Eider, Somateria mollissima, populations nesting in Arctic 
Canada. We surveyed Common Eiders in the Digges Sound region, Nunavut, on 19-20 July 1999, for comparison with 
similar surveys in 1980-1983. On six islands for which data were available in both periods, there were 4-5 times as many 
nests in 1999 as in the early 1980s. The mean clutch size late in incubation in 1999 (3.3 eggs per nest) was similar to that in 
1983 (3.2 eggs), but lower than in 1982 (3.9 eggs); timing of hatching was similar in the two periods. All three females 
caught during incubation were of the subspecies S. m. borealis. In contrast to Common Eiders 5. m. sedentaria nesting in 
the Belcher Islands, Common Eiders nesting in the Digges Sound region appear to be faring well. 

Key Words: Common Eider Somateria mollissima, clutch size, Digges Sound, egg size, populations, timing of hatching. 



Common Eiders, Somateria mollissima, are an 
important subsistence and cultural resource for Inuit 
people in Canada's eastern Arctic, in additon to being 
important components of arctic marine ecosystems 
(Reed 1986). Despite this, there is little information 
available on status and trends of arctic-nesting 
Common Eider populations. Historical data exist for 
only a few locations in eastern Arctic Canada: the 
Belcher Islands and nearby Quebec coast 
(Nakashima and Murray 1988), Ungava Bay 
(Chapdelaine et al. 1986), the West Foxe Islands 
(Cooch 1977; Cooch 1986), the Hell's Gate- 
Cardigan Strait region (Prach et al. 1986), and the 
Digges Sound region (Gaston et al. 1985). There is 
evidence that Common Eider, S. m. sedentaria, pop- 
ulations nesting in the Belcher Islands have declined 
sharply (Robertson and Gilchrist 1998). There also is 
evidence of long-term declines in S. m. v-nigra pop- 
ulations of northern Alaska and the western 
Canadian arctic (Suydam et al. 2000). Updated infor- 
mation on other populations is urgently required in 
order to assess their status and devise effective con- 
servation strategies (compare Reed and Erskine 
1986; Dickson 1997). 

We counted Common Eider nests on six islands 
in the Digges Sound region, Nunavut, on 19-20 
July 1999. Lying at the western end of Hudson 
Strait, Digges Sound appears to be near the south- 
ern limit of the breeding range of the migratory S. 
m. borealis subspecies, and near the northern limit 
of the breeding range of the non-migratory S. m. 
sedentaria subspecies (Abraham and Finney 1986). 
Common Eiders are an important source of meat, 
eggs, and down for people from the community of 



Ivujivik, Quebec, which lies in the heart of the 
region (Gaston et al. 1985). In this paper, we report 
the results of our 1999 surveys and make compar- 
isons with observations from 1980-1983, to assess 
the status of the Common Eider population nesting 
in this region. 

Study Area 

The Digges Sound region, and its seabird commu- 
nity, was described in detail by Gaston et al. (1985; 
see Figure 1). It is dominated by a very large aggre- 
gation (300 000 pairs) of Thick-billed Murres, Uria 
lomvia, that breed on the tall cliffs of East Digges 
Island and the Quebec mainland south and west of 
Cape Wolstenholme. There are also many small, 
low-lying islands that support nesting colonies of 
Common Eiders and other marine birds. Potential 
predators on Common Eider eggs and ducklings that 
we saw on these small islands included Herring 
Gulls, Larus argentatus. Glaucous Gulls, L. hyper- 
boreus, and Great Black-backed Gulls, L. marinus. 
The former two species breed in colonies throughout 
Digges Sound (note that there were >31 adult 
Glaucous Gulls with > 6 ca. 15 to 20 day old young 
on South Skerry #3 on 19 July in 1999; this colony 
was not reported in Gaston et al. 1985). Only one 
Great Black-backed Gull, a first-summer bird, was 
seen in the Digges Sound region during 1980-1983, 
but there were three adult-plumaged birds near South 
Skerry #1 on 19 July in 1999; we saw no sign of 
breeding. There was no evidence of predation by 
Arctic Foxes, Alopex lagopus, on any islands that we 
visited. A single Polar Bear, Vrsus maritimus, was 
present in Digges Sound in mid- July in 1999, but we 



22 



A 



2002 



HiPFNER, Gilchrist, Gaston, and Cairns: Common Eiders in Digges Sound 



23 




GINGI ISLAND 



Figure 1. Map of the southern Digges Sound region showing the islands that were surveyed for Common Eiders in 1999. 



saw nothing to suggest that it had been on any 
islands that we visited. 

Methods 

We surveyed six islands in the Nuvuk Islands 
region on 19 July 1999, including some of those on 
which the highest numbers of nests were found dur- 
ing surveys in the 1980s (in particular, the South 
Skerries; see Table 1, Figure 1). Results of our sur- 
veys should be comparable to those in the 1980s, 
because similar techniques were used; the one 
exception was that the number of nests was only 
estimated, not counted, on South Skerry #1 in 1981 
(see Table 1). In 1999, each island was surveyed on 
foot by a crew of 3 who walked 5 m apart. As the 
islands were too large to be covered in one sweep, 
the crew member at the outside of the line marked 
the extent of each pass with rocks. The crew then 
turned around and returned, counting nests on the 
opposite side of the line. In this manner, islands were 
completely surveyed with little risk of nests being 
missed or counted more than once. A nest was 
counted if it had a well-defmed cup with down; in 
this region, down is placed in the nest bowl after lay- 
ing has been initiated (H. G. Gilchrist unpublished), 
so that downy nest cups without eggs represent nest- 
ing attempts that have failed. When a nest was 
found, its contents (e.g., empty, two eggs, three eggs 



and one duckling, etc.) were called out to one crew 
member who recorded the information. We thus 
obtained information on clutch sizes, and a crude 
indication of the timing of hatching. We also mea- 
sured the length and maximum breadth of 100 eggs 
in 36 clutches. 

We attempted to survey island groups to the north 
on 20 July, but rough seas and thick fog hampered 
our efforts. We returned to South Skerry #1 late in 
the afternoon on 20 July, in order to catch adult 
eiders. Bill measurements and blood samples were 



Table 1. Common Eider nest counts in the Digges Sound 
region in different years. 







>'ear 




Island 


1981 


1983 


1999' 


South Skerry #1 


75^ 


42 


326(255) 


South Skerry #2 


- 


19 


66 (48) 


Pitsulak City 


3 


- 


2(2) 


Green 


4 


20 


25 (24) 


Yellow 


1 


- 


2(2) 


Black 





- 






'Counts in 1999 are: total nests with well-defined cups 
(total nests with > 1 egg/duckling); no such distinction was 
made on counts in 1981 or 1983. 
-Estimate only, no systematic count was made. 



24 



The Canadian Field-Naturalist 



Vol. 116 



taken to determine the subspecific status of these 
birds (see Mendall 1986). 

Results 

We caught three incubating female Common 
Eiders on South Skerry #1 on 20 July. Bill measure- 
ments indicated that all three were of the subspecies 
S. m. borealis (cf. Table 1 in Mendall 1986): frontal 
extensions ranged from 13.2 to 16.3 mm, nostril 
extensions 25.7 to 31.0 mm, and total bill lengths 
63.2 to 66.6 mm. 

On the six islands we surveyed in 1999, we count- 
ed 421 nests with well-defmed cups, of which 331 
had at least one egg/duckling (Table 1). Our total 
count in 1999 was about 4-5 times higher than 
counts on the same six islands in the early 1980s 
(Table 1). Note that the count of 158 nests (range: 
143-181) reported for the South Skerries in Gaston 
et al. (1985) was for all four of these islands, with 
the highest count from South Skerry #4 (73 nests). 
We did not do counts on South Skerry #3 or #4 in 
1999. 

In nests with at least one egg/duckling on South 
Skerry #1, South Skerry #2, and Green Island 
(n = 327; Table 1), clutch sizes late in incubation 
ranged up to 6 eggs, with a mean of 3.33 ± 1.03 (SD) 
eggs per nest. Common Eider clutch sizes varied 
among years in the Digges Sound region (ANOVA, 
^2 442= '7-53, P < 0.001). Tukey's tests (at the 
a = 0.05 significance level) indicated that the mean 
clutch size in 1999 was similar to that in 59 nests on 
the same three islands late in incubation in 1983 
(3.20 ± 1.13), but smaller than that in 59 nests late in 
incubation on Eider Island and the North Skerries in 
1982 (3.86 ± 1.09). The mean clutch size in 1999 
was similar to that reported for S. m. borealis popu- 
lations elsewhere in Canada (means of 2.72-3.52 
eggs; cf. Table 3 in Prach et al. 1986). 

At least one duckling was present in 0.9% of 
active nests (n = 331) found on 19 July, all of them 
on South Skerry #1 and #2. We also saw five broods 
on the sea off South Skerry #1 on this date, with 2-5 
duckhngs per brood. These observations suggest that 
hatching began about 17 July 1999, similar to the 
timing in the early 1980s: Common Eider ducklings 
were first seen on 17 July 1980, 21 July 1981, 14 
July 1982, and 20 July 1983. 

For 100 eggs from 36 entire clutches measured on 
South Skerry #1, Green Island, and Pitsulak City, the 
mean length was 75.4 + 2.7 (SD) mm (range: 
68.5-82.8 mm), and the mean breadth 49.3 ± 1.3 
(SD) mm (range: 46.8-53.3 mm). Eggs were not 
measured on surveys in the early 1980s, but 
Common Eider eggs in the Digges Sound region in 
1999 were similar in size to those laid by S. m. bore- 
alis elsewhere in Canada (mean lengths: 74.0 ± 3.3 
to 75.8 ±3.3 mm; mean breadths: 49.6 ± 1.7 to 
49.7 ± 1 .4 mm; cf. Table 2 in Prach et al. 1986). 



Discussion 

The total number of Common Eider nests that we 
counted on six islands in the Digges Sound region in 
1999 was about 4-5 times the number counted on 
the same six islands in the early 1980s. It can be dif- 
ficult to interpret the results of surveys such as these 
(Robertson and Gilchrist 1998). Many Common 
Eiders may forego breeding in years when environ- 
mental conditions are unfavourable (Coulson 1984), 
and as a result, low nest counts in such years could 
be wrongly interpreted as being indicative of popula- 
tion declines. Similarly, surveys that detect more 
nests, as did ours, could be wrongly interpreted as 
indicating population increases if the original sur- 
veys were carried out in unfavourable years with 
extensive non-breeding. While the latter is possible, 
we do not believe that this was the case here. 

For Common Eiders, breeding parameters such as 
egg size, clutch size, and the timing of laying are 
sensitive to variation in environmental conditions 
during the pre-laying period (Robertson 1995). Egg 
and clutch sizes that we measured were very similar 
to those reported for Common Eiders elsewhere in 
the eastern Canadian arctic, and clutch sizes were 
similar to, or smaller than, those in the Digges Sound 
region in the early 1980s. Moreover, the timing of 
hatching and dates of observations of first broods 
appeared to be very similar during the two survey 
periods. Collectively, these results suggest that it is 
unlikely that conditions were so poor that only low 
proportions of eiders bred in all years when surveys 
were carried out in the early 1980s. Given that sur- 
vey techniques were similar in the two time periods, 
involving counts of all active and failed nests, we 
conclude that the substantial increase in the number 
of Common Eider nests in the Digges Sound region 
probably reflects an actual increase in the size of the 
local breeding population. 

If so, this raises two important questions. First, 
what factors might have caused the increases in the 
numbers of Common Eiders nesting in this region 
since the early 1980s? And second, if real, are 
increases in this region indicative of increases in 
Common Eider S. m. borealis populations elsewhere 
in eastern Arctic Canada? 

We do not have definitive answers to either ques- 
tion. Given that there were no obvious differences in 
environmental conditions between the two survey 
periods, the dramatic increases that we detected may 
have been due to lower levels of local human 
exploitation than occurred previously. Common 
Eider populations in the Foxe Islands declined dra- 
matically between 1956 and 1976, apparently due to 
local harvesting practices. Many eider colonies in 
that region are easily reached by hunters traveling by 
boat from Cape Dorset, and the subsistence harvest 
of eggs, and especially the shooting of females, had 
dramatic effects on the population (Cooch 1986). It 



I 



2002 



HiPFNER, Gilchrist, Gaston, and Cairns: Common Eiders in Digges Sound 



25 



may be noteworthy that the largest increases in the 
number of Common Eider nests in the Digges Sound 
region occurred on the South Skerries. These islands 
are off the regular travel routes, and rarely are visited 
by people from Ivujivik (Gaston et al. 1985). As a 
result, they may function as safe nesting refuges dur- 
ing times when few people are venturing out specifi- 
cally to harvest eiders or their eggs. Unfortunately, 
no data exist to allow us to directly examine the pos- 
sibility that the apparent increase was due to lower 
local harvests of Common Eiders, or eider products, 
in recent years. We also have no information with 
which to assess the role that immigration from local 
or more distant colonies might have played. 

Might the apparent increases that we detected in 
Common Eider populations in the Digges Sound 
region reflect increases elsewhere? Although popula- 
tions of S. m. sedentaria appear to have declined 
considerably in the Belcher Islands, an island group 
to the south of Digges Sound, this was probably due 
to severe ice conditions in Hudson Bay that caused 
high overwinter adult mortality in this non-migrator>' 
subspecies during the early 1990s (Robertson and 
Gilchrist 1998). Causes of declines in populations of 
S. m. \-nigra in northern Alaska and the western 
Canadian arctic are unknown (Suydam et al. 2000). 

It is both costly and logistically difficult to con- 
duct surveys of Common Eiders at arctic colonies, 
and they cannot normally be carried out on a large 
geographic scale. Instead, small clusters of nesting 
islands are usually surveyed and the results cautious- 
ly extrapolated to the larger population (Cooch 
1986). However, surveys conducted in regions where 
eiders have historically been, or are curtently being, 
subjected to local subsistence harvests cannot reli- 
ably be extrapolated to non-harvested populations, 
because local anthropogenic effects could easily 
swamp large-scale environmental effects. Conse- 
quently, repeat surveys of colonies elsewhere are 
now required to assess trends in S. m. borealis popu- 
lations in other areas. Given these considerations, it 
is apparent that an ideal monitoring scheme for 
Common Eiders would include both harvested and 
non-harvested populations. Although this would add 
substantially to the cost of monitoring programs, it 
should allow investigators to distinguish between the 
local effects of human harvesting and environmental 
effects that operate over a broader geographic scale. 
This would aid greatly in management. 

Acknowledgments 

We thank David and John Geale. as well as the 
residents of Ivujivik, Quebec, for assistance with the 
surveys. The surveys were supported by the 
Canadian Wildhfe Service. Memorial University of 
Newfoundland, the Natural Sciences and Engineer- 
ing Research Council of Canada, the Northern 
Scientific Training Program, and the Polar Conti- 



nental Shelf Project of Natural Resources Canada. 
We thank Scott GilHland for insightful comments on 
a previous draft of this manuscript. 

Literature Cited 

Abraham, K. F., and G. H. Finney. 1986. Eiders of the 
eastern Canadian Arctic. Pages 55-73 m Eider Ducks in 
Canada. Edited by A. Reed. Canadian Wildlife Service 
Report Series Number 47. Ottawa. 

Chapdelaine, G., A. Bourget, W. B. Kemp, D. J. 
Nakashima, and D. J. Murray. 1986. Population 
d'Eider a duvet pres des cotes du Quebec septentrional. 
Canadian Wildlife Service Report Series 47: 39-50. 

Cooch, F. G. 1977. Changes in the avifauna of the West 
Foxe Islands, Northwest Territories, 1956-1976. Cana- 
dian Field-Naturalist 91: 314-317. 

Cooch, F. G. 1986. The numbers of nesting Northern 
Eiders on the West Foxe Islands, NWT. in 1956 and 1976. 
Canadian Wildlife Service Report Series 47: 1 14— 1 18. 

Coulson, J. C. 1984. The population dynamics of the 
Eider Duck Somateria mollissima and evidence of exten- 
sive non-breeding by adult ducks. Ibis 126: 525-543. 

Dickson, D. L. 1997. King and Common eiders of the 
western Canadian Arctic. Canadian Wildlife Service 
Occasional Paper Number 94. Ottawa. 

Gaston, A. J., D. K. Cairns, R. D. Elliot, and D. G. 
Noble. 1985. A natural history of Digges Sound. 
Canadian Wildlife Service Report Series Number 46, 
Ottawa. 

Mendall, H. L. 1986. Identification of eastern races of the 
Common Eider. Canadian Wildlife Ser\ice Report Series 
47: 82-88. 

Nakashima, D. J., and D. J. Murray. 1988. The Com- 
mon Eider {Somateria mollissima sedentaria) of eastern 
Hudson Bay: a survey of nest colonies and Inuit ecologi- 
cal knowledge. Environmental Studies Revolving Funds 
Report Number 102. Ottawa. 

Prach, R. W., A. R. Smith, and A. Dzubin. 1986. 
Nesting of the Common Eider near Hell's Gate-Cardigan 
Strait polynya, 1980-1981. Canadian Wildlife Service 
Report Series 47: 127-137. 

Reed, A. 1986. Eiderdown harvesting and other uses of 
Common Eiders in spring and summer. Canadian Wild- 
life Ser\ice Report Series 47: 138-146. 

Reed, A., and A. J. Erskine. 1986. Populations of the 
Common Eider in eastern North America: their size and 
status. Canadian Wildlife Service Report Series 47: 
156-162. 

Robertson, G. J. 1995. Annual varation in common eider 
Qgg size: effects of temperature, clutch size, laying date, 
and laying sequence. Canadian Journal of Zoology 73: 
1579-1587. 

Robertson, G. J., and H. G. Gilchrist. 1998. Evidence of 
population declines among Common Eiders breeding in 
the Belcher Islands. Northwest Territories. Arctic 51: 
375-381. 

Suydam, R. S., D. L. Dickson, J. B. Fadely, and L. T. 
Quackenbush. 2000. Population declines of King and 
Common Eiders of the Beaufort Sea. Condor 102: 
219-222. 

Received 26 April 2000 
Accepted 28 March 2002 



Flathead Chubs, Platygobio gracilis, in the Upper Missouri River: 
The Biology of a Species at Risk in an Endangered Habitat 

Shannon J. Fishery, David W. Willis^, Michael M. Olson^, and Steven C. Krentz^ 

'Minnesota Department of Natural Resources, Windom, Minnesota 56101 USA 

^Corresponding author. Department of Wildlife and Fisheries Sciences, South Dakota State University, Brookings, South 

Dakota 57007 USA.; e-mail: david_willis@sdstate.edu 
^U.S. Fish and Wildlife Service, Bismarck, North Dakota 58501 USA 

Fisher, Shannon J., David W. Willis, Michael M. Olson, and Steven C. Krentz. 2002. Flathead Chubs, Platygobio gracilis, 
in the upper Missouri River: The biology of a species at risk in an endangered habitat. Canadian Field-Naturalist 
116(1): 26-41. 

Flathead Chub, Platygobio gracilis, populations have declined in portions of their range where natural flood-pulses and 
functioning backwaters have been eliminated. Observations of these declines have led ecologists to consider both the 
species and their at-risk habitats. Backwater and sandbar habitats in the Missouri River, North Dakota were sampled during 
a high flow (1997) and an average flow (1999) year. Of the 817 larval fishes captured in the backwaters, none were 
Flathead Chubs. Significantly fewer (P<0.01) Flathead Chub adults and juveniles were captured in the backwaters than 
from sandbar habitats during all sample periods. Male chubs were all mature at 110-mm total length (TL) and age 2, 
whereas females were not all mature until age 3 and 170-mm TL. In 1997, Ostracoda, Hemiptera, and Copepoda dominated 
the diets, whereas in 1999, Coleoptera, Trichoptera, and Hymenoptera were utilized in significantly greater amounts 
(P<0.05). Flathead Chub maximum age was 5 and the mean back-calculated total lengths at age were 104-, 153-, 186-, and 
223-mm TL for ages 1-4. Incremental growth analyses indicated that greater increases in total length were attained in 1997 
than in 1999; however, body condition was significantly higher in 1999 (P< 0.006). The contradiction between growth and 
condition cannot be explained; however, it could be energetically advantageous to maintain a more fusiform body shape 
during high flows. Although the backwaters were not important physical habitat for Flathead Chubs, backwater productivi- 
ty may have critically contributed to the prey base during the high flow period of 1997. 

Key Words: Flathead Chub, Platygobio gracilis, Cyprinidae, habitat alterations, food habits, Coleoptera, Copepoda, back- 
waters, Missouri River, North Dakota. 



Cross et al. (1986) and Pfleiger and Grace (1987) 
reported that Flathead Chub, Platygobio gracilis 
Richardson, populations have decreased by as much 
as 98% over portions of their historical range. Grady 
and Milligan (1998) and Gelwicks et al. (1996) both 
observed Flathead Chub declines in the channelized 
and impounded segments of the Missouri River. 
These declines are not well understood; however, 
degraded habitats are a suspected cause (Lee et al. 
1980). Researchers have correlated sensitive species 
declines with channel and flow modifications (Hesse 
et al. 1989). In addition to the problems of popula- 
tion loss and habitat modifications, Flathead Chub 
are difficult to study because their life history has 
not been thoroughly documented. 

Fisher (1999) noted that a segment of the Missouri 
River in North Dakota contained Flathead Chubs and 
was maintaining historical habitat characteristics, 
including natural flood-pulse patterns from 
Yellowstone River inflow and a functioning lateral 
relationship with floodplain backwaters. Scott and 
Nielsen (1989) and Sabo and Kelso (1991) demon- 
strated that backwater habitats are important rearing 
and nursery areas for numerous native fishes. 
Additionally, Amoros (1991) suggested that flood- 
plain flushing transports invertebrates from backwa- 
ters to the main channel, providing an important 



food resource for native fishes obligated to flowing 
water habitats. Eckblad et al. (1984) observed sub- 
stantial increases in channel invertebrate densities 
below connection points with floodplain wetlands 
and Kennedy (1979) suggested that restricted back- 
water discharge may create invertebrate voids in the 
main channel. Therefore, understanding the relation- 
ships that Flathead Chub populations maintain with 
their habitats and food resources under varying water 
conditions may be critical. 

The relatively unknown status and function of 
Flathead Chub populations in North American river 
systems remain a point of interest. Furthermore, 
opportunities to study Flathead Chubs and other 
native fishes in relatively unaltered habitats and 
hydrographs have become increasingly rare. There- 
fore, the objectives of this study were to (1) assess 
general Flathead Chub biology, including age and 
growth, food habits, habitat use, size structure, and 
maturation patterns, (2) contrast life history patterns 
between two years with substantially different 
hydrographs, and (3) assess and discuss the potential 
importance of backwater habitats to Flathead Chubs. 
This reference information from a stable Flathead 
Chub population in a healthy river stretch should 
help biologists evaluate populations in degraded lotic 
systems. 



26 



I 



2002 



Fisher, Willis, Olson, and Krentz: Flathead Chubs in Upper Missouri 



27 



Study Area 

The Missouri River study area lies between the 
Yellowstone River confluence and Lake Sakakawea 
in North Dakota (48°N, 103.9°W; river kilometers 
2510-2518; river mile markers 1569-1574). This 
Missouri River stretch was selected because the 
unregulated Yellowstone River discharge provides a 
nearly historic flood-pulse hydrograph. Also within 
this stretch, several functioning backwater wetlands 
and numerous clusters of sandbar habitats are pre- 
sent. We monitored two backwaters and two sandbar 
complexes for use by Flathead Chubs. The two back- 
waters were described as off-channel habitats that 
connected to the Missouri River during one or more 
periods annually and maintained minimal flow 
(Fisher 1999). Surface areas for both backwaters 
were variable depending upon the prevailing hydro- 
logic conditions; however, each was characterized 
by annual inundation of terrestrial vegetation, silt 
and mud dominated substrate, presence of marsh 
smartweed, Polygonum coccineum, and depths that 
generally did not exceed 1.5 m. The sandbar com- 
plexes were each located in close proximity to the 
outlet channels of the above-mentioned backwaters 
and also varied in extent and persistence due to 
hydrologic conditions. Although the location and 
extent of sandbars was dynamic, the habitat was 
dominated by sand substrates, tended to be free from 
woody or other vegetative debris, and ranged in 
depth from 0.2 to 2.0 m. 

Methods 

Sampling Schedule and Habitat Analyses 

To provide a distinct comparison, we sampled 
Flathead Chubs in the Missouri River during one 
hydrograph cycle that approximated mean flow 
(1999) and during another cycle that included sub- 
stantially greater flow volume (1997; USGS 1999*; 
Figure 1). In 1997 and 1999, temperature, turbidity, 
and dissolved oxygen data were collected from chan- 
nel-sandbar and backwater habitats. See Table 1 for 
sampling schedule information. We were unable to 
sample sandbars during sample period 2 in both 
1997 and 1999 due to high waters from mountain 



30000 




/\ 




«■ 25000 




A 


1997 


(A 

•^ 20000 




J \k 






JM v\ 




$ 15000 
O 




r^iiiL^ 




U- 10000 


mJ^\. J 


/ llllllllll^^ 




5000 


i i|liiilliiiiiiiiiii^^ 




iiiilliiiiiiiaiiiiiiiiiiiiiiiiiiiiii 


30000 ■ 








g" 25000 ■ 






1999 


JO 

■V 20000 ■ 








■ — • 








5 15000 ■ 








"- 10000 ■ 


V"^ /-^ 


^ ^%.. 




5000 ■ 


„^- — -^ 




"^^-^--^..^-^^ 


90 120 


150 180 210 240 






Day of year 





Figure 1 . Cumulative daily flow (m^/sec) for the Missouri 
River below the Yellowstone River confluence and 
above the Lake Sakakawea headwaters in North 
Dakota for calender days 75-260 of 1997 and 1999. 
The solid line represents the total sum of the flow 
estimates for gauging stations in the Missouri River 
and Yellowstone River upstream from the conflu- 
ence. The shaded area represents the 40-year mean 
flow for the period 1959-1999. 



snowmelt that inundated the habitat from mid-May 
through mid- June. Data were tested for normality 
with a Shapiro-Wilkes test (UNIVARIATE proce- 
dure; SAS 1990). Likely due to small sample size, 
not variance, none of these data sets were normally 
distributed; therefore, a nonparametric Kruskal- 
Wallis analysis of variance (AOV) by ranks 
(NPARIWAY procedure; SAS 1990) was used to 
assess main effects. When a significant difference 
was detected, a Mann- Whitney U test (NPARIWAY 
procedure; SAS 1990) was used to locate those dif- 
ferences. Statistical significance for these and all 
other analyses was set at a= 0.05. 

Fish Sampling and Analyses 

Fish were collected during each sample period in 
the backwaters and from sandbar habitats with a bag 
seine (30-m long X 1.8-m high, 1.8-m' bag. 6-mm bar 



Table 1. Sampling schedule used in 1997 and 1999 in the study area, including the Missouri 
River main channel (CH), sandbar complexes (SB), and associated backwaters (BW). 
Habitats sampled during each sample period are denoted with an X. Some data, due to high 
flows and minimal habitat presence, were not available (NA). 





Calender Days 


Month 




Habitats Sampled 




Sample Period 


CH 


SB 


BW 


1 


114-120 


April 


X 


X 


X 


2 


139-145 


May 


X 


NA 


X 


3 


183-189 


July 


X 


X 


X 


4 


225-23 1 


August 


X 


X 


X 


5 


255-261 


September 


X 


X 


X 



28 



The Canadian Field-Naturalist 



Vol. 116 



mesh). Six seine hauls were completed in each habitat 
type per sample period between 1000 and 1700 hours. 
The seine was pulled perpendicular to the backwater 
shorelines or with the current along sandbar margins 
for approximately 20 m and then arched to the shore- 
line. Captured fishes from the backwater samples 
were counted, measured to the nearest mm total length 
(TL), and released. Fishes captured during sandbar 
seining were also counted and measured; however, up 
to 30 Rathead Chubs from each of the following total 
length groups per sample period were euthanized and 
returned to the laboratory for biological analyses. The 
total length groups were < 60, 60-99, 100-139, 
140-179, and > 179 mm. 

The Flathead Chub relative abundance was ex- 
pressed as number/seine haul and length-frequency 
histograms were created. Relative abundance data 
were tested for normality. Data not normally dis- 
tributed were transformed [loglO(n+l)] and again 
tested for normality. The relative abundance data 
were found to not be normally distributed, even after 
transformation; therefore, we used a nonparametric 
Mann-Whitney U test on the untransformed data to 
assess the differences between habitat types among 
sample periods within each year. The same analysis 
was used to evaluate differences between habitats 
during each sample period between years. To pro- 
vide additional samples for biological assessments in 
the two length groups exceeding 140-mm TL, a gill 
net (21-m long X 2-m high with one 7-m panel each 
of 1-, 2-, and 3-cm bar measure mesh) was drifted 
for short distances on the deeper portions 
(1.5-2.0 m) of the sandbar flats. Gill-net captured 
Flathead Chubs were added to samples for biological 
assessments; however, these data were not included 
in catch-per-unit-effort (CPUE) analyses. 

To better assess the potential utility of backwaters 
as nursery and rearing habitats for Flathead Chubs 
during their early life history, larval fishes were col- 
lected with modified quatrefoil light traps (Floyd et 
al. 1984; 25-cm high X 30-cm wide with two 2-mm 
and two 4-mm slot openings). Ten randomly placed 
light trap sets were completed in each backwater 
during each sample period. Attempts to sample lar- 
vae in calm channel areas with the light traps failed 
due to physical trap damage and sedimentation. The 
light traps were deployed between 1600 and 1800h 
and emptied prior to 1 100 h the following day. Two 
12-h photochemical light sticks were used as the 
light source in each light trap each night. 
Photochemical light stick intensity and light duration 
varies with temperature; however, Kissick (1993) 
found that photochemical sticks attract larvae for at 
least 1 h. All photochemical sticks used in this study 
were found to continue glowing up to 24 h after ini- 
tial use; regardless, light sticks were replaced each 
night. Larval fishes were preserved in 5% formalin 
and returned to the laboratory for identification. Case 



specimens were sent to the Larval Fish Laboratory at 
Colorado State University for positive identification. 
Once identified, the specimens were enumerated and 
indexed as number/trap night. 

General Biological Evaluation 

In the laboratory, euthanized Flathead Chubs were 
again measured to the nearest mm TL and weighed 
to the nearest 0. 1 g. Additionally, scales and otoliths 
were removed from September-sampled fish for age 
and growth analyses, stomachs were removed for a 
seasonal food habits assessment, and gonads of each 
fish were inspected to determine sex and maturity 
stage. Log 10 transformed weight-length regressions 
were completed for each year and sample period 
(REG procedure; SAS 1990) and analyzed with an 
analysis of covariance (GLM procedure; SAS 1990) 
to determine if intercepts and slopes differed 
between years. 

Age and Growth Assessment 

We attempted to age the September-captured 
Flathead Chubs using both otoliths and scales. Scales 
were aged directly from a projected microfiche 
image. The focus, annuli, and radius data from each 
scale were digitized into the DISBCAL software 
program and mean length at age was determined 
using the Fraser-Lee method (Frie 1982). A regres- 
sion analysis of scale radius on total Flathead Chub 
length was conducted within DISBCAL to determine 
the best available intercept, or a- value, for the back- 
calculations. Otoliths were prepared for reading 
using both vertical cross-sectioning and horizontal 
grinding to help expose the otolith circuli; however, 
the annuli were not discemable. Therefore, all of the 
age and growth analyses were based on scale annuli. 
Mean length at age for Flathead Chubs from both 
years combined was determined to provide a general 
description of growth at this geographic location. 

To assess differences in growth between the high- 
flow year of 1997 and the mean-flow year of 1999, 
the mean growth increments for 2-cm TL Flathead 
Chub subgroups were determined for the first 258 
days of each respective year. Martyn and Schmulbach 
(1978) suggested that annulus formation did not occur 
in Flathead Chubs until late May; therefore, the 
growth increment from the time of capture to the last 
annulus may only represent growth for 100-1 15 days. 
Using an AOV for unbalanced data sets, we complet- 
ed a two-way AOV with year and length category as 
the main effects (GLM procedure; SAS 1990). The 
interaction term between the main effects was not sta- 
tistically significant (P = 0.156); therefore, we com- 
pleted a one-way AOV between years within each 
length category to determine if the Flathead Chubs 
accumulated total length at a greater rate in 1997 or 
1999. We also completed an AOV among length cate- 
gories within each year to determine if different length 
groups accumulated total length at significantly differ- 
ent rates. When a significant difference was detected. 



2002 



Fisher, Willis, Olson, and Krentz: Flathead Chubs in Upper Missouri 



29 



we used a Tukey multiple-range test to identify those 
differences (TUKEY option in GLM procedure; SAS 
1990). 

Food Habits Assessment 

Stomach contents from the euthanized Flathead 
Chubs were enumerated. Food habits were assessed 
for feeding uniformity by determining the frequency 
of occurrence, while feeding intensity on each prey 
type was investigated by determining the percent by 
number for each prey item within each Flathead Chub 
length group. Frequency of occurrence was the pro- 
portion of individuals in each Flathead Chub length 
group that contained the prey item. The percent by 
number for each prey item was determined for stom- 
ach contents for each fish that contained food and the 
mean values for each length group were reported. 
Bo wen (1996) noted that proportional diet data are not 
normally distributed; therefore, a Mann- Whitney U 
test was used to detect significant proportional differ- 
ences between years within each sample period and 
Flathead Chub length group. Because our objective 
was to assess food habit differences between different 
hydrologic conditions, no comparison among length 
groups within each year was completed. 

Results and Discussion 

Habitat Comparison 

Channel habitat parameters, including tempera- 



ture, dissolved oxygen, and turbidity were not signif- 
icantly different (P>0.05) from sandbar measure- 
ments in any collection period during either 
collection year (Table 2); therefore, the results and 
discussion will focus on the comparisons of backwa- 
ters with sandbar habitats only. Backwater and sand- 
bar temperamres were more similar in 1997 than in 
1999. Backwater temperatures in 1997 were 0.4 to 
1 .4°C warmer than the sandbar habitats, but were not 
significantly different during any months (P > 0.05). 
In 1999, backwater temperatures were significantly 
higher (P<0.05) in all months by 1.8 to 4.5°C 
(Table 2). September was the only sample period in 
1997 with significantly higher backwater dissolved 
oxygen levels (P< 0.05); whereas in 1999, July was 
the only sample period without significantly greater 
backwater dissolved oxygen concentrations. 
Turbidity levels were significantly lower (P < 0.05) 
in the backwaters during all sample periods in 1997 
and 1999 (Table 2). 

Sandbar habitat temperatures during the July and 
September sample periods of 1997 were significantly 
warmer (P<0.05) than in 1999 by 1.6 to 1.7°C. In 
contrast, the backwaters were significantly cooler 
during May and September of 1997 (P<0.05) than 
the same sample periods of 1999 by 4.1 and 2.4°C, 
respectively (Table 2). During July, August, and 
September of 1999 in both sandbar and backwater 
habitats, dissolved oxygen levels were significantly 



Table 2. Mean temperature, dissolved oxygen, and turbidity data collected in the Missouri River, North Dakota, in 1997 
and 1999. Data were collected at designated time periods in backwaters (BW), the main channel (CH), and near sandbar 
complexes (SB). Standard errors are noted in parentheses. The results of three separate statistical comparisons are 
described in this table. Within each year and month, the three habitats were compared and the statistical results are coded 
with the letters a, b, and c. Within each year and habitat type, the sample periods (months) are compared and the statistical 
results are coded with the letters m, n, o, and p. Within each habitat type and month, the 1997 and 1999 statistical compari- 
son is coded with the letters x and y. Within each set of comparisons, means with the same letters were not statistically dif- 
ferent (P> 0.05). Each mean was derived from five measurements. 







1997 






1999 


■ 


Parameter/Period 


BW 


CH 


SB 


BW 


CH 


SB 


Temperature (°C) 














April 


amxll 7 


amxl] 2 


amxn.3 


amxl 2.6 


bmxlO.6 


►^"'"10.8 


May 


anxl4_2 


amnxl3 6 


NA 


"nyl8.3 


bnyl6.1 


NA 


July 


^0^22. 8 


apy21.3 


aoy21.4 


ao''22.8 


boxl9 4 


b«xi9.8 


August 


aox2L9 


aoxl9 9 


aoxJ9 9 


"0"23.3 


box 19.8 


b«"20.0 


September 


an" 16.7 


any 16.0 


any 16.3 


any 19.1 


bnxl4 4 


bnxl4.6 


Dissolved oxygen (mg/L) 














April 


"""9.1 (0.3) 


"""8.5 (0.2) 


"""8.6 (0.3) 


"ny9.4 (0.2) 


'^n"8.3(0.1) 


^^"'"8.5 (0.2) 


May 


""y9.3 (0.3) 


"py9.6 (0.2) 


NA 


""»7.8 (0.3) 


b"'"6.6 (0. 1 ) 


NA 


July 


'"""7.0 (0.2) 


"n«6.7(0.1) 


""^"6.8 (0.2) 


"■"y8.4 (0.3) 


"ny8.5(0.1) 


"n'y8.4 (0.2) 


August 


^■""7.7 (0.3) 


""i""7.2 (0.2) 


"ni""7.2 (0.3) 


"ny9.3 (0.2) 


Kv8.3(0.1) 


'^"'>8.1 (0.2) 


September 


^'""8.3 (0.1) 


hn"7.7(0.1) 


•^""7.7(0.1) 


"»yll.0(1.0) 


t'"y9.2(0.1) 


►'"y9.3 (0.2) 


Turbidity (NTU) 














April 


"""66(18) 


bny94 (7) 


bny99 (5) 


anx46 (11) 


bnx74 (9) 


•^""82 (3) 


May 


amxl7(8) 


hpy216(24) 


NA 


•'""33 (10) 


'^p^l35(2) 


NA 


July 


"'^"16(3) 


h"yl56(6) 


boy 160 (6) 


""'"10(3) 


'^""112(3) 


•^'^16(5) 


August 


"-""14(5) 


h"y85 (8) 


►'"y89 (3) 


""'"8 ( 1 ) 


>^""65 (3) 


•^""72 (4) * 


September 


an,x 1 1 (2) 


h"«38 (5) 


h"'"39 (5) 


•"""9 (2) 


brm4() (3) 


hinx^g (4) ■ 



30 



The Canadian Field-Naturalist 



Vol. 116 



higher than in 1997 (P<0.05; Table 2). Turbidity 
levels were not significantly (P > 0.05) different in 
the backwater habitats between 1997 and 1999, even 
though sandbar turbidity levels were significantly 
higher (P< 0.05) in April, July, and August of 1997 
(Table 2). 

In 1997, the high flow created more floodplain 
connection points with longer duration. As a result, 
the 1997 temperature and dissolved oxygen levels in 
the backwaters were quite similar with those found 
in the channel. In 1999, these same variables operat- 
ed more independently with less influence from 
channel connections (Table 2). Regardless of year, 
the backwaters maintained the ability to filter chan- 
nel connection inflow and without exception, back- 
waters had significantly lower turbidity levels 
(P< 0.01; Table 2). This filtering capability, warmer 
water temperatures, an abundance of potential food 
resources such as zooplankton and benthic inverte- 
brates (Fisher 1999), and more static conditions 
would appear to create conditions sought and pre- 
ferred by some native fishes as nursery, rearing, 
feeding, and refuge habitats. 

Fish Sampling 

In 1997 and 1999 combined, 817 larval fish were 
captured in the light traps; however, none were veri- 
fied as Flathead Chubs. Of these larvae, 70% were 
Ictiobus spp. (Buffalo fishes), 12% were Common 
Carp, Cyprinus carpio, and 8% were River Carp- 
sucker, Carpiodes carpio. Additionally, only eight 
Flathead Chub juveniles and adults (>3-cm TL) 
were captured with the bag seine in the backwaters 
(Table 3). Significantly more Flathead Chubs, 
> 12/seine haul, were captured in the sandbar habitats 
during all sample periods and both years (P<0.01; 
Table 3). 

Although greater numbers of Flathead Chubs were 
captured on sandbar habitats, there also existed sev- 
eral significant differences in mean CPUE among 
sample periods. During both years, greater numbers 
of Flathead Chubs could be sampled near sandbar 



complexes later in the year (Table 3). In addition to 
higher CPUE, more large Flathead Chubs, particular- 
ly those exceeding 16-cm TL, could be captured dur- 
ing August and September, with the greatest range in 
size structure occurring in September during both 
1997 and 1999 (Figure 2). 

Even though numerous larval fishes were sampled 
in the backwater habitats, no chubs of any genera 
were captured in the light traps. If the chub larvae 
were present, they were not sampled, possibly due to 
negative phototaxic habits or data collection timing 
problems. Likewise, the low CPUE indicated mini- 
mal presence of juvenile and adult Flathead Chubs in 
the backwater habitat. We hypothesized that high 
backwater connectivity to the channel would allow 
increased Flathead Chub use of the backwater habi- 
tats. Even in the presence of near record connectivity 
during 1997, Flathead Chub use of the backwaters as 
physical habitat was limited. 

The increased presence of large (> 16-cm TL) 
Flathead Chubs on the sandbars in the September 
samples (Figure 2) might suggest that the primary 
breeding adults stage, spawn, and recover in other 
channel habitats, as their conspicuous absence dur- 
ing the entire season except for the fall samples was 
easily noted. Peters and Holland (1994) found that 
all life stages of Flathead Chubs in the Platte River, 
Nebraska, utilized channel regions with sandy sub- 
strates and depths < 50 cm. The sandbar habitats 
appeared to be important feeding and rearing areas 
for juvenile Flathead Chubs during all sample peri- 
ods. In addition, the age-0 Flathead Chubs, repre- 
sented by newly recruited individuals <4-cm TL, 
appeared on the sandbar habitats in late summer and 
early fall. The observation of age-0 Flathead Chub 
using the sandbar complexes could be an important 
life history documentation. 

Sexual Maturity 

Gonadal assessment in September-captured 
Flathead Chubs revealed that some males began to 
exhibit maturing testes by 60-mm TL, but likely did 



Table 3. Catch per unit effort for Flathead Chub captured with bag seines (number/seine haul) in two upper 
Missouri River backwaters (BW) and on two sandbar complexes (SB). The standard error of each mean is noted 
in parentheses. There are two sets of statistical comparisons denoted in this table. Within each habitat type and 
month, the letters a and b represent the comparisons between the 1997 and 1999 data points. Within each habitat 
and year (i.e., each column of means), the letters x and y represent the comparison among the five sample peri- 
ods (months). For each set of comparisons, means with the same letters are not statistically different (P>0.05). 
Each mean was derived from six measurements. 





1997 




1999 


Sample period 


BW 


SB 


BW 


SB 


April 

May 

July 

August 

September 


^^0.2(0.1) 
"0.2 (0.2) 
^'^0.3 (0.2) 
^''O.O (0.0) 
^"^0.0 (0.0) 


t^M2.3(3.2) 

NA 
'"'y24.2(3.1) 
>'y37.0(13.3) 

''''y29.2 (9.7) 


^"0.2(0.1) 
'^0.3 (0.2) 
ax0.2(0.1) 
^0.0 (0.0) 
^'^0.0 (0.0) 


•"'18.7(8.2) 

NA 

bM 2.5 (6.3) 

''y52.0(11.5) 

''y55.8 (22.5) 



2002 



Fisher, We.lis, Olson, and Krentz: Flathead Chubs in Upper Missouri 



31 



25 
20 



52 



^ 39 



> » <i) 



3 3 



5- 26 



o 



3 0) 

3 C 
< S> 



o 



o 



13 



70 
56 
42 
28 



14 



32 



24 



E o 



iJ 3 16 



8 




8 12 16 20 24 



8 12 16 20 24 
Total length (cm) 



Total length (cm) 

Figure 2. Length-frequency histograms of Flathead Chub captured from sandbar habitats in the Missouri River. North 
Dakota during four sample periods in 1997 and 1999. The frequencies represent the total catch from six seine hauls 
per sample period. 



not spawn until the following season. Of male 
Flathead Chubs captured in April, 66% had devel- 
oped testes at 75-mm TL. At 1 10-mm TL, 100% of 
male Flathead Chubs had reached sexual maturity. 
The smallest Flathead Chub that contained devel- 
oped ovaries in April was 85-mm TL. Only 21% of 
the chubs that appeared to be female and were less 
than 125-mm TL contained mature eggs. By 155- 
mm TL, 80% of the female Flathead Chubs were 



mature, but not until 170-mm TL was 100% female 
sexual maturity noted. 

Olund and Cross (1961) suggested that sexual 
maturity was attained by both sexes at about 85-mm 
standard length in Alberta, and Brown (1971) noted 
well-developed gonads in Flathead Chubs approxi- 
mately 82-mm TL in Montana. Conversely, Gould 
(1985) found that the minimum total length for male 
and female maturation in the Musselshell River. 



32 



The Canadian Field-Naturalist 



Vol. 116 



Table 4. Frequency of occurrence (%) for stomach contents of Flathead Chubs collected in 1997 and 1999 from sandbar 
habitats in the Missouri River, North Dakota. Sample periods, total number (N), and percent of empty stomachs (E) are 
noted. Food items included Ostracoda (OST), Diptera (DIP), Ephemeroptera (EPH), Hemiptera (HEM), Hymenoptera 
(HYM), Odonata (ODO), Orthoptera (ORT), Plecoptera (PLE), Trichoptera (TRI), Copepoda (COP), Coleoptera (COL), 
and macrophytes (MAC). Total length groups (TL; mm), included 1=<60, 2=60-99, 3=100-139, 4=140-179, and 5=>179. 
Absent groups imply that no specimens were collected. 



Month 


Year 


Length 


N 


E 


OST 


DIP 


EPH 


HEM 


HYM 


ODO 


ORT 


PLE 


TRI 


COP 


COL 


MAC 


April 


97 


1 


8 


0.0 


100.0 


12.5 


0.0 


25.0 


0.0 


0.0 


0.0 


0.0 


12.5 


50.0 


0.0 


0.0 


April 


99 


1 


13 


0.0 


16.4 


23.1 


0.0 


0.0 


7.7 


0.0 


0.0 


0.0 


0.0 


15.4 


92.3 


0.0 


April 


97 


2 


30 


46.7 


36.7 


26.7 


0.0 


20.0 


0.0 


3.3 


0.0 


0.0 


16.7 


26.7 


0.0 


0.0 


April 


99 


2 


30 


13.3 


3.3 


40.0 


3.3 


9.9 


0.0 


0.0 


0.0 


13.3 


3.3 


3.3 


76.7 


0.0 


April 


97 


3 


13 


0.0 


69.2 


30.8 


0.0 


30.8 


0.0 


30.8 


0.0 


0.0 


53.8 


38.5 


0.0 


0.0 


April 


99 


3 


29 


10.3 


6.9 


55.2 


3.4 


10.3 


17.2 


17.2 


0.0 


13.8 


27.6 


3.4 


72.4 


0.0 


April 


97 


4 


4 





25.0 


50.0 


0.0 


0.0 


0.0 


25.0 


25.0 


0.0 


75.0 


75.0 


0.0 


0.0 


April 


99 


4 


10 


0.0 


0.0 


20.0 


20.0 


40.0 


10.0 


10.0 


0.0 


50.0 


30.0 


0.0 


80.0 


0.0 


July 


97 


1 


17 


17.6 


52.9 


5.9 


0.0 


29.4 


0.0 


0.0 


0.0 


0.0 


11.8 


41.2 


0.0 


0.0 


July 


99 


I 


30 


30.0 


0.0 


13.3 


0.0 


0.0 


0.0 


0.0 


0.0 


0.0 


43.3 


3.3 


26.7 


0.0 


July 


97 


2 


30 


50.0 


36.7 


13.3 


0.0 


20.0 


20.0 


0.0 


0.00 


0.0 


6.7 


16.7 


0.0 


0.0 


July 


99 


2 


29 


13.8 


0.0 


6.9 


0.0 


0.0 


0.0 


0.0 


0.0 


79.3 


3.4 


0.0 


0.0 


0.0 


July 


97 


3 


10 


0.0 


50.0 


50.0 


10.0 


30.0 


0.0 


0.0 


50.0 


0.0 


10.0 


30.0 


0.0 


0.0 


July 


99 


3 


30 


0.0 


0.0 


16.7 


0.0 


0.0 


0.0 


0.0 


0.0 


6.7 


93.3 


3.3 


6.7 


0.0 


July 


97 


4 


3 


0.0 


66.6 


0.0 


33.3 


33.3 


0.0 


0.0 


66.6 


0.0 


66.6 


33.3 


0.0 


0.0 


July 


99 


4 


14 


14.3 


0.0 


0.0 


0.0 


0.0 


0.0 


0.0 


0.0 


0.0 


78.6 


14.3 


7.1 


0.0 


July 


97 


5 


2 


0.0 


50.0 


50.0 


0.0 


50.0 


0.0 


0.0 


100.0 


0.0 


0.0 


0.0 


0.0 


0.0 


July 


99 


5 


3 


33.3 


0.0 


0.0 


0.0 


0.0 


0.0 


0.0 


0.0 


0.0 


33.3 


33.3 


0.0 


0.0 


August 


97 


1 


9 


0.0 


55.6 


33.3 


0.0 


44.4 


0.0 


0.0 


0.0 


0.0 


33.3 


33.3 


0.0 


0.0 


August 


99 


1 


1 


0.0 


0.0 


100.0 


0.0 


0.0 


0.0 


0.0 


0.0 


0.0 


0.0 


0.0 


10.0 


0.0 


August 


97 


2 


30 


53.3 


33.3 


3.3 


13.2 


19.8 


0.0 


3.3 


3.3 


0.0 


3.3 


23.1 


0.0 


0.0 


August 


99 


2 


30 


6.7 


0.0 


50.0 


46.7 


0.0 


0.0 


0.0 


0.0 


6.7 


13.4 


3.3 


10.0 


0.0 


August 


97 


3 


13 


0.0 


53.8 


7.7 


23.1 


53.8 


0.0 


38.5 


46.2 


0.0 


30.8 


30.8 


0.0 


0.0 


August 


99 


3 


27 


11.1 


0.0 


40.7 


85.2 


0.0 


0.0 


0.0 


0.0 


0.0 


11.1 


0.0 


7.4 


0.0 


August 


97 


4 


4 


0.0 


75.0 


0.0 


50.0 


50.0 


0.0 


0.0 


50.0 


0.0 


50.0 


0.0 


0.0 


0.0 


August 


99 


4 


30 


6.7 


0.0 


40.0 


73.3 


0.0 


3.3 


3.3 


0.0 


6.7 


30.0 


0.0 


10.0 


0.0 


August 


99 


5 


4 


0.0 


0.0 


50.0 100.0 


0.0 


0.0 


0.0 


0.0 


50.0 


50.0 


0.0 


0.0 


0.0 


Sept 


97 


1 


30 


16.7 


50.0 


30.0 


0.0 


46.7 


0.0 


0.0 


0.0 


0.0 


20.0 


30.0 


0.0 


0.0 


Sept 


99 


I 


13 


30.8 


0.0 


15.4 


0.0 


0.0 


53.8 


0.0 


0.0 


0.0 


7.7 


0.0 


15.4 


0.0 


Sept 


97 


2 


30 


6.7 


56.7 


33.3 


53.3 


53.3 


6.7 


29.7 


6.7 


3.3 


19.8 


26.4 


0.0 


0.0 


Sept 


99 


2 


30 


46.7 


0.0 


16.7 


0.0 


0.0 


43.3 


0.0 


0.0 


0.0 


0.0 


0.0 


23.3 


3.3 


Sept 


97 


3 


20 


0.0 


70.0 


25.0 


35.0 


50.0 


10.0 


20.0 


25.0 


5.0 


20.0 


40.0 


0.0 


0.0 


Sept 


99 


3 


28 


60.7 


0.0 


7.1 


0.0 


0.0 


17.9 


0.0 


0.0 


0.0 


10.7 


0.0 


0.0 


3.6 


Sept 


97 


4 


13 


0.0 


46.1 


53.8 


15.4 


30.8 


0.0 


0.0 


46.1 


0.0 


46.1 


23.1 


0.0 


0.0 


Sept 


99 


4 


23 


34.8 


0.0 


17.4 


0.0 


4.3 


39.1 


0.0 


0.0 


0.0 


0.0 


0.0 


17.4 


13.0 


Sept 


97 


5 


12 


0.0 


33.3 


41.7 


50.0 


25.0 


8.3 


0.0 


41.7 


8.3 


25.0 


16.7 


0.0 


0.0 


Sept 


99 


5 


10 


70.0 


0.0 


10.0 


0.0 


0.0 


10.0 


0.0 


0.0 


0.0 


0.0 


0.0 


0.0 


20.0 



Montana, was 113 and 123 mm, respectively. Al- 
though there is some disagreement about length at 
maturity, our results suggest that some males are 
reproductively active at age 1 and are all mature by 
age 2, whereas only a very few females are mature at 
age 1 and not until age 3 have all females entered the 
breeding population. Scarnecchia et al. (2000) also 
noted mature males at age 1 and a few mature 
females at age 2 in the Yellowstone River, Montana. 
Our observations are similar to Bishop (1975) and 
Martyn and Schmulbach (1978), who suggested that 
sexual maturity was attained by age 2 and 105-mm 
standard length for the species as a whole. 

Food Habits 

In 1997, the most frequently consumed prey 



across all length groups of Flathead Chubs during 
the April and July samples were Ostracoda, 
Copepoda, Hemiptera (dominated by Corixidae), 
and Diptera. During the late summer and into 
autumn, Ostracoda, Hemiptera, and Copepoda con- 
tinued to demonstrate frequent presence in the stom- 
ach contents; however, Ephemeroptera and 
Trichoptera became more prevalent in the diets 
(Table 4). The 1999 frequency of occurrence data 
indicated that fewer Flathead Chubs were consum- 
ing Copepoda and Hemiptera. In addition, 
Ostracoda were nearly absent from the diets. 
Alternatively in 1999, Flathead Chub feeding strate- 
gies appeared to be more focused on Coleoptera in 
the spring and early summer and Hymenoptera and 



I 



2002 



Fisher, Willis, Olson, and Krentz: Flathead Chubs in Upper Missouri 



33 




<60 60-99 99-139 140-179 



<60 60-99 99-139 140-179 



Total length group (mm) Total length group (mm) 

Figure 3. Numerical proportions (%) of primary prey taxa consumed by Flathead Chubs collected in April of 1997 and 
1999 from the Missouri River, North Dakota. For each pair of bars (1997 and 1999), a statistical comparison was 
conducted. Those pairs with a * designation were significantly different (P<0.05). 



macrophytic plant seeds in the late summer and 
autumn (Table 4). 

Numerically, Ostracoda dominated the Flathead 
Chub diets in 1997, with Copepoda, Hemiptera, 
Trichoptera, and Diptera also contributing (Figures 
3-6). In 1999, Ostracoda, Hemiptera, and Copepoda 
were significantly less abundant in the Flathead 
Chub diets (P < 0.05) during numerous sample peri- 
ods and among most length groups (Figures 3-6). 
The significant decline in several prey taxa was off- 
set by significant increases in the numeric propor- 
tions of Coleoptera, Diptera, Trichoptera, Ephemer- 
optera, and Hymenoptera during several seasons and 
for various length groups (all tests were P<0.()5; 
Figures 3-6). 

The food habits assessment provided evidence 
that the diets of Flathead Chubs in the high water 
year of 1997 were substantially different than those 



observed in the typical water year of 1999. Olund 
and Cross (1961) noted that Flathead Chubs col- 
lected in Alberta rivers had diets similar to those 
reported here, primarily consuming Corixidae, 
Hymenoptera, Diptera, and Coleoptera. In addition. 
Brown (1971) suggested that terrestrial insects 
were an important food resource during portions of 
the year. 

Cellot and Bournard (1987) noted that backwaters 
produce a number of invertebrates relatively uncom- 
mon in channel habitats. Some of those backwater 
taxa were the primary contents noted in our Flathead 
Chub diet analysis in 1997. such as Copepoda zoo- 
plankton and Hemiptera. Although Ostracoda can be 
abundant in the backwater habitats, their origin is not 
clear. Pennak (1989) noted that seed shrimps 
(Ostracoda) are well-equipped to thrive in lotic and 
lentic habitats. The Corixidae and Copepoda, howev- 



34 



The Canadian Field-Naturalist 



Vol. 116 



c 
o 

r 
o 
a 
o 



c 
o 

r 
o 

a 
o 



o 

r 
o 

a 
o 



1997 
1999 



Ostracods 




Diptera 



i£ 



ii 



Trichoptera 

*I ^ •T 



•I 



Coleoptera 



^ 



Total length group (mm) 



Total length group (mm) 



Figure 4. Numerical proportions (%) of primary prey taxa consumed by Flathead Chubs collected in July of 1997 and 
1999 from the Missouri River, North Dakota. For each pair of bars (1997 and 1999), a statistical comparison was 
conducted. Those pairs with a * designation were significantly different (P < 0.05). 



er, both tend to be confined to lentic waters and 
exhibit low densities in lotic habitats (Merrit and 
Cummins 1984; Pennak 1989). Therefore, the evi- 
dence suggests that the high flows in 1997 likely 
contributed to the altered diets by transporting prey 
taxa from the floodplain to the channel and/or by 
washing in-channel food resources away from sand- 
bars at a greater rate than during a typical flow year. 
In 1999, the consumption of Coleoptera during 
April and extending into the other sample months 
was pronounced. Although reported as Coleoptera, 
nearly all of the beetles observed in the Flathead 
Chub stomachs belonged to the family Cicindelidae 
(tiger beetles). Borror and White (1970) noted that 
tiger beetles, where present, tended to be very abun- 
dant and inhabited open shorelines and beaches. 
Numerous tiger beetles were observed on the sand- 



bars, and Borror and White (1970) also suggested 
that the larvae tend to burrow into sandy areas dur- 
ing their juvenile stages. We can only speculate 
about the mechanisms that made tiger beetles avail- 
able to Flathead Chubs; however, during the spring 
and summer of 1999, many tiger beetles were 
observed in the drift and it is possible that as sand- 
bars shift, late-stage larvae and possibly adults are 
expunged from their burrows and become more sus- 
ceptible to fish predation. Alternatively, Flathead 
Chubs may actively forage into the substrates. Live 
Flathead Chubs held in the laboratory demonstrated 
digging behaviors while foraging, at times burying 
themselves past their pectoral fins. In 1997, the 
absence of tiger beetles from the diet was conspicu- 
ous. It is possible that the inundation of sandbar 
habitats, which occurred earlier than normal in 1997, 






2002 



Fisher, Willis, Olson, and Krentz: Flathead Chubs in Upper Missouri 



35 



c 
o 

o 

a 
o 



c 
o 

-E 
o 
a 
o 



c 
o 

o 

a 
o 



100 
80 
60 
40 
20 

98 
75 
60 
45 
30 
15 


24 ■ 

18 ■ 

12 

6 \ 





• 






Diptera 




1 1 ' 


■ 




T^ 




i 








^ 


T 


T 



1997 
1999 



Ephemeroptera 

•I 




Total length group (mm) 



Total length group (mm) 



Figure 5. Numerical proportions (%) of primary prey taxa consumed by Flathead Chubs collected in August of 1997 and 
1999 from the Missouri River, North Dakota. For each pair of bars (1997 and 1999), a statistical comparison was 
conducted. Those pairs with a * designation were significantly different (P < 0.05). 



removed the staging substrates, either repelling the 
beetles away from the river bottom in search of bet- 
ter habitats or transporting the organisms out of the 
feeding area. 

The consumption of backwater-originated prey 
resources by Flathead Chubs during 1999, and more 
extensively in 1997, suggests that backwater prey 
production could be important to Flathead Chubs in 
the Missouri River, particularly during high-flow 
periods. In a relatively backwater-poor section of the 
Yellowstone River, Montana, upstream from our 
study site, Scarnecchia et al. (2000) also collected 
Flathead Chub in 1997, but found that 98% of the 
stomachs were empty. The high 1997 flow rates may 
have transported more typical Flathead Chub prey 
into unusable areas; therefore, the increased con- 
sumption of backwater-oriented food resources may 
have been a compensatory effort to obtain nutrition. 



We did not document any empty stomach propor- 
tions greater than 70%, and for most Flathead Chub 
length categories and sample periods, empty stom- 
ach proportions were typically < 40% (Table 4). It is 
unclear if the backwater prey availability was suffi- 
cient to provide adequate rations for the Flathead 
Chub population. 

Age and Growth 

To complete the back-calculation process, we 
needed to obtain a Flathead Chub intercept, or a- 
value, for the scale radius to total body length rela- 
tionship. A regression analysis based on all fish col- 
lected provided an a-value of 32 mm. This value 
seemed somewhat high considering that many small- 
er species, such as the darters, Etlwostoma spp., typi- 
cally have a-values less than 22 mm (Carlandcr 
1997) and larger species such as Walleye, Stizo- 
stedion vitrcum and White Bass, Moronc chrysops. 



36 



The Canadian Field-Naturalist 



Vol. 116 



c 
o 

-E 
o 

a 
o 



o 
o 

a 
o 



c 
o 

o 

a 
o 



1997 
1999 



Hemiptera 




Total length group (mm) 



Total length group (mm) 



Figure 6. Numerical proportions (%) of primary prey taxa consumed by Flathead Chubs collected in September of 1997 
and 1999 from the Missouri River, North Dakota. For each pair of bars (1997 and 1999), a statistical comparison 
was conducted. Those pairs with a * designation were significantly different (P < 0.05). 



have a-values greater than 30 mm (Carlander 1982; 
Beck et al. 1997). To help validate the use of a 32- 
mm intercept during the back-calculations, we fur- 
ther assessed the Flathead Chub specimens in our 
collection that were <40-mm TL and in good body 
condition. Upon further inspection, all of the 
Flathead Chubs < 30-mm TL were scaleless and did 
not appear to be physically damaged in any way to 
indicate that scales may have been dislodged during 
handling. Of the remaining Flathead Chubs < 40-mm 
TL, 45% of those between 30- and 35-mm TL were 
scaled and by 38-mm TL, all specimens had scales 
with discernable circuli. Therefore, the a-value of 
32 mm was used with relative confidence. 

Aging the Flathead Chub scales was a difficult pro- 
cess and the agreement proportion between the two 
initial readers (both inexperienced) was 42%. 
Likewise, Scarnecchia et al. (2000) indicated that 
Flathead Chub scale aging was challenging. Gould 



(1985) relied upon length-frequency histograms to 
identify year classes after unsuccessful attempts to age 
Flathead Chubs with scales, vertebrae, and opercula. 
The length-frequency histograms that we formulated 
for this Missouri River population did not allow us to 
confidently identify cohorts (Figure 2). Although 
early age groups can be discerned on the length-fre- 
quency histograms, variability was too high to rely on 
that technique. Therefore, we consulted other cyprinid 
researchers and aging experts. After receiving a 
cyprinid aging tutorial, we re-aged the scales with 
three readers (2 inexperienced and 1 experienced) and 
reached a consensus age on 96% of the scales. The 
remaining 4% were eliminated from the analysis due 
to our inability to assign an age. Our age analysis indi- 
cated that the maximum age of Flathead Chub in our 
collection was 5 years. Therefore, considering growth 
and maturation patterns, many females may only 
release eggs in one or two seasons. 






2002 



Fisher, Willis, Olson, and Krentz: Flathead Chubs in Upper Missouri 



37 



Table 5. Mean back-calculated total lengths (TL; mm) at age for Flathead Chubs collected 
in 1997 and 1999 from the Missouri (MO) River in North Dakota. Standard errors are in 
parentheses. Also noted are total lengths (mm) at age for Flathead Chubs from Perry Creek, 
Iowa (Martyn and Schmulbach 1978) and from the Peace River, Alberta [Bishop (1975); fork 
lengths converted to total length using conversion in Martyn and Schmulbach (1978)]. 



Age 



MO River (N) 



MO River (TL) 



Perry Creek 



Peace River 



1 


85 


104.1 (2.8) 


77.8 (0.6) 


100.4 (NA) 


2 


37 


152.9 (4.7) 


109.6 (0.5) 


123.6 (NA) 


3 


17 


186.4 (3.8) 


130.0 (0.5) 


152.2 (NA) 


4 


6 


223.2 (6.9) 


148.0 (2.3) 


164.9 (NA) 


5 


1 


267.0 (NA) 


NA 


179.2 (NA) 



The back-calculated lengths at the time of annulus 
formation for each age group suggested that Flathead 
Chubs, on average, increase their length by approxi- 
mately 46% between ages 1 and 2, but then growth 
slows and plateaus at annual length increases of 
19-22% during the remainder of their life (Table 5). 



Incremental growth analyses for Flathead Chubs col- 
lected in September of each year indicated that por- 
tions of the Flathead Chub length range, including 
those between the total lengths of 95 and 135 mm, 
significantly increased at a more rapid rate during 
1997 than did similar sized fish in 1999 (P<0.02; 






O 

u 
ox 

c 



50 -r 



45 ■ 



40 ■ 



S 35 



30 



S 25 



20 




1%-^' 96-^^' ^X^Al^* ^^SA^* ^65^1^ ^^%,\9^ .\9^ 



Total length group (mm) 



Figure 7. Mean growth increments for Flathead Chubs collected in the Missouri River, North Dakota in September of 1997 
and 1999. The increments represent estimated growth from the time of annulus fonnation to the time of capture 
each year. The letters a and b represent the statistical comparison between years for each total length group noted 
on the X-axis (vertical comparison). The letters x, y, and z denote the comparison results among total length groups 
within each sample year (horizontal comparison). Means with the same letters were not statistically different 
(P > 0.05). 



38 



The Canadian Field-Naturalist 



Vol. 116 



Figure 7). With the exception of those age-1 chubs 
less than 95-mm TL, all length groups had higher 
mean increments in 1997 than in 1999; however, dif- 
ferences for groups >134-mm TL were not statisti- 
cally significant (P<0.05). In 1997, the longest 
growth increments were attained by chubs between 
95 and 174 mm, whereas in 1999, chubs between 95 
and 135 mm had significantly lower growth incre- 
ments (P < 0.05; Figure 7). 

As compared with the maximum age of 5 reported 
here, Martyn and Schmulbach (1978) found Flathead 
Chubs up to age 4 in Perry Creek, Iowa; however, 
Scarnecchia et al. (2000) suggested that Flathead 
Chubs in the Yellowstone River, Montana, reached 
age 7 and Bishop (1975) noted that the maximum 
life span of Flathead Chubs in the Peace River, 
Canada was 10 years. Additionally, the longest 
Flathead Chub in our sample was 275-mm TL, 
whereas Kristensen (1980) reported capturing 293 
Flathead Chubs in the Peace-Athabasca River delta 
in Canada that had a mean TL of 345 mm. The sub- 
stantially higher total length and longevity reported 
for Flathead Chubs in the northern portions of the 
species range may be attributed to less disruption of 
recruitment patterns and cooler water temperatures 
that moderate metabolic activity, ultimately extend- 
ing the average potential life span. The population of 
Flathead Chub utilizing the sandbar habitats in this 
study was dominated by individuals < age 3. 

Brown (1971) reported lengths at ages 1-3 for 
Flathead Chubs in Montana; however, no indication 
was given pertaining to how these lengths and ages 
were derived or if the measurements were in total, 
fork, or standard length (Table 5). Martyn and 
Schmulbach (1978) aged Flathead Chub scales col- 
lected from specimens in Perry Creek, a Missouri 
River tributary in Iowa, and noted lengths consider- 
ably shorter for each age group than those reported 
in our study (Table 5). Gould (1985) also noted three 
Flathead Chub total length modes at 43-, 81-, and 
116-mm TL in the March length-frequency his- 
togram from the Musselshell River, Montana, but 
did not suggest that the modes represented age 
groups. Our assigned ages and back-calculated 
lengths at age differed from several other reports, 
but were similar to those reported at ages 1-5 in the 



Peace River, Alberta (Bishop 1975). We believe that 
our aging was done appropriately and given the wide 
variability of habitats and phenotypical expression 
across the species range, our results should be repre- 
sentative of the Flathead Chub population found in 
the study area. 

Although understanding mean growth patterns for 
Flathead Chubs as a population is valuable, we were 
particularly interested in the growth rates during 
1997 and 1999 when the diets significantly differed. 
The cohorts of Flathead Chubs in 1997 utilized prey 
that were likely backwater originated, including 
Copepoda and Hemiptera, at a significantly higher 
rate than the same cohorts did in 1999 (Figures 3-6). 
Food resources that appeared to have backwater 
origination were nearly absent from the chub diets in 
1999. This dietary absence may have influenced the 
growth rates of different length groups to varying 
degrees, especially when considering feeding effi- 
ciency and food item assimilation patterns that can 
cause substantial changes in the bioenergetics of dif- 
ferent cohorts within a species (Adams and Breck 
1990). 

In addition to changes in diet, environmental con- 
ditions can also alter bioenergetic functions and 
affect fish growth rates and body condition by 
directly influencing metabolism and prey dynamics 
(e.g.. Kitchen et al. 1977; Swenson 1977). In 1997, 
water temperatures were cooler in the early portions 
of the annual flood pulse (i.e., April and May), but 
were significantly warmer (P<0.05) from July 
through September (Table 2). The higher summer 
and fall temperatures in 1997 may be attributable to 
the extended exchange of water between the flood- 
plain and the channel. Waters retained in backwaters 
and other floodplain wetlands tend to be substantial- 
ly warmer, as was demonstrated in 1999 when the 
backwaters functioned in an uncoupled manner from 
the channel (Table 2). The higher temperatures in 
1997 may have contributed to the increases in 
growth increments that could also be noted on the 
scales from fish collected in 1999, particularly those 
that were ages 1 and 2 in 1997. 

Body Condition 

Regardless of sample period, Flathead Chubs cap- 
tured in the mean flow year of 1999 weighed less at 



I 



Table 6. Weight-length equations for Flathead Chubs captured in various sample periods in 1997 and 1999 from the 
Missouri River in North Dakota. Statistical comparisons (analyses of covariance), represented by their p-values, between 
years for intercepts (a) and slopes (b) are noted. Weight (wt) is in grams and total length (TL) is in mm. 



Weight-length equations 



P-values 



Period 



1997 



1999 



April log,„wt = -5.369+3.()65(log„)TL) 

July log,(,wt= -5.62 1+3. 226(log,„TL) 

August log|„wt = -4.902+2.853(log|„TL) 

September log,„wt = -5.349+3.063(logn,TL) 



log,„wt = -4.680+2.802(log„TL) 
log,„wt = -5.028+2.997(log„)TL) 
log,(,wt = -5.245+3. 107(log,JL) 
log wt = -4.91 8+2.938(]ogTL) 



0.001 


0.001 


0.001 


0.001 


0.006 


0.001 


0.001 


0.001 



2002 



Fisher, Willis, Olson, and Krentz: Flathead Chubs in Upper Missouri 



39 




01) 



2.0 

1.5 



-^ 1 

OX) 

-2 0.0 



-0.5 ■ 



April 




July 






2.0 • 


-So 


1.5 ' 


-^ 




Si 

0£ 


1.0 ■ 






o> 




o 


0.5 ■ 


^^ 




&£ 




^ 


0.0 ■ 




-0.5 ■ 




1.7 1.8 1.9 2.0 2.1 2.2 2.3 1.4 1.6 1.8 2.0 2.2 2.4 2.6 
logjQtotal length (mm) logjototal length (mm) 

Figure 8. Weight-length relationships (log 10 transformed) from Flathead Chubs collected in the Missouri River, North 
Dakota, during various periods of 1997 and 1999. The cross-hair markers denote chubs collected in 1997 and the 
circles represent those collected in 1999. The regression relationship for each set of data is depicted with a solid 
line. Statistical analyses for these data are summarized in Table 6. 



all comparable lengths than in 1997 (Figure 8). The 
intercept values and slopes of these regression analy- 
ses were all significantly different between years 
during each sample period (P< 0.006; Table 6). 
During some sample periods, the 1999 predicted 
weights for some Flathead Chub lengths exceeded 
the 1997 predicted weights by as much as 50%. 

The weight-length equations reported here were 
similar to those documented by Gould (1985); how- 
ever, that study did not identify any significant dif- 
ferences in slopes or intercepts among sample peri- 
ods. It would be reasonable to hypothesize that 
when the environment favors increased growth rates 
(based on total length), that the habitats would also 
be favorable for individuals to maintain high body 
condition (plumpness). Although the logic of this 



argument seems apparent, our data suggest that 
Flathead Chubs in 1997 acquired greater total length 
additions, but did so while they were less plump. 
Fish condition has been correlated with prey avail- 
ability in many lentic populations (Wege and 
Anderson 1978; Liao et al. 1995; Marwitz and 
Hubert 1997; and Porath and Peters 1997), but we 
are aware of no such analyses for lotic fish popula- 
tions. 

We cannot explain the contradictory condition and 
growth observations discussed above; however, it 
could be hypothesized that during a year when flow 
velocity is high (e.g., 1997) it is ecologically advan- 
tageous for a fish to be more fusiform than when 
under conditions of lower flow rates. Fisher et al. 
(1996) noted that healthy burbot. Lola lota, popula- 



40 



The Canadian Field-Naturalist 



Vol. 116 



tions from lotic systems consistently maintained 
lower overall body condition; however, annual varia- 
tion in these condition measures was not assessed. 
Although Flathead Chubs in 1997 were thinner and 
had to cope with higher turbidity levels (Table 2), 
the population was also provided with food 
resources and warmer water temperatures due to the 
higher rate of floodplain interaction. Scott and 
Crossman (1973) noted considerable morphological 
variation in Flathead Chubs across its North 
American range and Olund and Cross (1961) sug- 
gested that subspecies, based on morphological fea- 
tures that included differences in total length to cir- 
cumference ratios, be recognized. However, Bailey 
and Allum (1962) hypothesized that the manifesta- 
tions were strictly related to environmental condi- 
tions, not genetic strains. The substantial differences 
that we observed between two years within the same 
Flathead Chub population support the plasticity sug- 
gestion of Bailey and Allum (1962). 

Flathead Chub food habits are often referred to in 
general, but seldom supported with documentation 
(e.g., McLane 1978; Baxter and Stone 1995). Brown 
(1971) reported that biologists in Montana believe 
the Flathead Chubs feed on aquatic invertebrates, 
extensively utilize abundant terrestrial insects that 
become part of the drift, and are sufficiently omniv- 
orous to feed on vegetative matter. Our results sup- 
port these assumptions, including the sporadic feed- 
ing on plant tissues. Our data indicate that diets vary 
substantially during differential hydrograph cycles 
and, when combined with varied flows and other 
environmental conditions, influence growth rates 
and condition. The backwater-produced prey 
appeared to be utilized by small to mid-sized 
Flathead Chubs. We were unable to determine if 
Flathead Chubs prefer the backwater organisms or 
are using their availability to compensate for absent 
channel resources during high flow periods. 
Riverine ecologists (e.g., Kennedy 1979; Eckblad et 
al. 1984) have assumed that backwater habitats are 
critical to native fishes evolved in large river sys- 
tems. We documented limited direct physical use of 
backwater habitats by Flathead Chubs during any 
life history stage; however, the indirect benefits of 
floodplain prey production and water conditioning 
are possibly critical during high flow hydrographs. 

Acknowledgments 

We thank the U.S. Fish and Wildlife Service 
(Region 6), the South Dakota Cooperative Fish and 
Wildlife Research Unit, South Dakota State Uni- 
versity, and the U.S. Geological Survey Species at 
Risk program for funding and administering this 
research. We also express our appreciation to 
Charles Pyle, Randy Sheik, Steve Wilson, Nate 
Olson, Dan Moon, Ryan Doorenbos, Danielle 
Johnson, and Gene Galinat for their assistance with 
field data collection and laboratory analyses. We 



thank Mike Brown for statistical advice, several 
reviewers, and the North Dakota State Department of 
Fish and Game, particularly Fred Ryckman and Greg 
Power for their cooperation and logistical assistance. 
This manuscript has been approved for publication 
by the South Dakota Agricultural Experiment Station 
as Journal Series Number 3187. 

Documents Cited (marked * in text) 

USGS (U.S. Geological Survey). 1999. Water resources 
of Montana: historical water data files. http://waterdata. 
usgs.gov/nwis-w/mt. 

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Kanawha River, West Virginia. Journal of Fish Biology 
35: 21-27. 

Swenson, W. A. 1977. Food consumption of walleye 
(Stizostedion vitreum) and sauger (S. canadense) in rela- 
tion to food availability and physical conditions in Lake 
of the Woods, Minnesota. Shagwa Lake, and western 
Lake Superior. Journal of the Fisheries Research Board 
of Canada 34: 1643-1654. 

Wege, G. J., and R. O. Anderson. 1978. Relative weight 
(Wr): a new index of condition for largemouth bass. Pages 
79-91 in New approaches to the management of small 
impoundments. Edited by G. D. Novinger and J. D. 
Dillard. North Central Division, American Fisheries 
Society, Special Publication (5). Bethesda. Maryland. 

Received 1 May 2000 
Accepted 5 March 2002 



, 



Abundance and Distribution of Breeding Waterfowl in the 
Great Clay Belt of Northern Ontario 

R. Kenyon Rossi, Kenneth F. Abraham^, Ted R. Gadawski^, Robert S. Rempel^, 
T. Shane Gabor^, and Ron Maher^ 

'Environment Canada, Canadian Wildlife Service, Ontario Region, 49 Camelot Drive, Ottawa, Ontario KIA 0H3 Canada 

[Corresponding Author] 
^Ontario Ministry of Natural Resources, Science, Development and Transfer Branch, 300 Water Street, 3'^'* Floor N, 

Peterborough, Ontario K9J 8M5 Canada 
3R.R. # 2, Shanty Bay, Ontario LOL 2L0 Canada 
'^Ontario Ministry of Natural Resources, Centre for Northern Forest Ecosystems Research, Lakehead University Campus, 

Thunder Bay, Ontario P7B 5E1 Canada 
^Institute for Wetland and Waterfowl Research, c/o Ducks Unlimited Canada, Oak Hammock Marsh Conservation Centre, 

P.O. Box 1 160, Stonewall, Manitoba ROC 2Z0 Canada 
^Ducks Unlimited Canada, 614 Norris Court, Unit # 1, Kingston, Ontario K7P 2R9 Canada 

Ross, R. Kenyon, Kenneth F. Abraham, Ted R. Gadawski, Robert S. Rempel, T. Shane Gabor, and Ron Maher. 2002. 
Abundance and distribution of breeding waterfowl in the Great Clay Belt of northern Ontario. Canadian Field- 
NaturaUst 116(1): 42-50. 

The abundance and distribution of breeding waterfowl in the Great Clay Belt of northern Ontario was determined through 
helicopter surveys of 117 fixed plots (2 X2km each) during the nest initiation periods from 1988 to 1990. This area has 
higher fertility, flat topography, high water table and better access than the surrounding Boreal Forest, and therefore has 
greater potential for increased waterfowl production through habitat management. Overall breeding density averaged 1 12.5 
indicated breeding pairs per 100 km^, 68% being of the four most common species [Mallard (Anas platyrhynchos). Ring- 
necked Duck {Aythya collaris), American Black Duck {Anas rubripes), and Common Goldeneye (Bucephala clangula)]; 
13 other species were encountered. The average total of breeding waterfowl for the region was estimated at 59330 pairs. 
Distributions of the species were related to ecodistrict and to surficial geology. The more northerly of the two main ecodis- 
tricts had higher densities of American Black Ducks, Ring-necked Ducks, Common Goldeneyes, and Canada Geese 
(Branta canadensis). Mallard and Hooded Merganser (Lophodytes cucullatus) distributions correlated with presence of sur- 
ficial clay and moraines, respectively. Less common species including Green- winged Teal {Anas crecca) and American 
Wigeon {Anas americana) appeared to be concentrated in smaller-scaled habitat features (beaver pond sequences and estu- 
arine marshes, respectively). Results generally agreed with those of earlier Clay Belt surveys. Total breeding density of 
waterfowl is slightly higher than that of surrounding regions. 

Key Words: ducks, populations, habitat, boreal, forest, Ontario 

The boreal forest region in eastern Canada has methods were applied that breeding density esti- 

been largely ignored as an important area for water- mates for the full range of waterfowl species could 

fowl production owing to its relatively low fertility be made (Dennis 1974; Ross 1987). These intensive 

and often difficult accessibility when compared to surveys revealed that different physiographic units of 

the prairies. An initial assessment by Hanson et al. the boreal forest had characteristic waterfowl species 

(1949) based on the limited information available at compositions. One such unit is the Great Clay Belt, 

the time concluded that the area was of minimal also known as the Northern Clay Section (B4 in 

importance to most waterfowl populations. How- Rowe 1972), and hereafter termed the Clay Belt in 

ever, as more quantitative surveys were applied, ini- this paper. In Ontario, the Clay Belt extends east 

tially by the U.S. Fish and Wildlife Service in the from Hearst to the Quebec border and south approxi- 

1950s and 1960s (Addy et al. 1952, Kaczynski and mately to Timmins (see Figure 1); the remainder of 

Chamberlain 1968), it became clear that, while this unit occupies an area of roughly similar size in 

breeding densities were low, the vast area of the contiguous Quebec. The Clay Belt is unique in hav- 

boreal forest made its contribution to continental ing a rich clay soil in contrast to low fertility habitats 

waterfowl stocks substantial (Wellein and Lumsden of muskeg and exposed-bedrock shield surrounding 

1964). The accuracy of these earlier surveys was it. The higher fertility of the Clay Belt is evidenced 

limited by the difficulty of observing waterfowl in by higher rates of wetland occupancy by waterfowl 

forested wetlands from fixed-wing aircraft than in adjoining areas (Dennis 1974). Moreover, the 

(Chamberlain and Kaczynski 1965). In Ontario, it combination of its general fertility and flat topogra- 

was only when more intensive and efficient survey phy, high water table, and relative accessibility due 



42 



2002 Ross, Abraham, Gadawski, Rempel, Gabor, and Maher: Breeding Waterfowl 



43 



Human Geography 






Ecological Classification 

Lac Motagami Ecoregion 

■ Ecodistrict 29 
S Ecodistrict 30 
W Other Ecodistricts 

E3 James Plain Ecoregion 
Chapleau Plains Ecoregion 




Surficial Geology 

Lacustrine Sedinnents (Clay) 
Ground Moraine 
End Moraine 
Marine Sedinnents 



Figure 1. Geographical locations, ecological classification, and surficial geology units of the Great Clay Belt 
of Ontario. 



to the extensive network of roads for logging and 
mining, gives the Clay Belt greater potential for 
enhanced waterfowl production through habitat man- 
agement than in the neighbouring boreal forest. This 
importance is well-recognized and the Clay Belt is 
included as a priority area in the Eastern Habitat 
Joint Venture of the North American Waterfowl 
Management Plan (NAWMP Update 1994). 

Maximizing the effectiveness of management 
activities requires knowledge of the breeding distri- 
butions and habitat preferences of various waterfowl 
species frequenting the area. A study of waterfowl 
productivity across the Clay Belt of Ontario was 



therefore undertaken jointly by Ducks Unlimited 
Canada, the Ontario Ministry of Natural Resources 
and the Canadian Wildlife Service. The main objec- 
tives were to: (1) assess species composition, densi- 
ty, and distribution of the waterfowl community. (2) 
determine habitat correlates of nesting and brood- 
rearing waterfowl, and (3) develop a wildlife habitat 
map using LANDSAT. This paper describes water- 
fowl species composition, density and distribution 
across the Clay Belt, relates these to broad habitat 
patterns in the area, and estimates the contribution of 
the Clay Belt to overall waterfowl populations in 
northern Ontario. 



44 



The Canadian Field-Naturalist 



Vol. 116 



Methods 

Study Area 

The Clay Belt is a relatively flat, humid forest 
region corresponding to an area occupied by former 
glacial lakes Ojibway and Barlow (Rowe 1972; 
Sharpe and Brodie 1931). Bedrock is mostly overlain 
by sedimentary clays and extensive peatland. There 
are relatively few large lakes, and streams and rivers 
are generally low gradient compared to adjacent 
exposed Precambrian shield areas. Tree species of 
the Clay Belt reflect this topography and geological 
history. The main tree species is Black Spruce 
(Picea mariana) which occurs on flat lowlands and 
moister uplands. In areas of better local drainage, 
species include White Spruce {P. glauca). Trembling 
Aspen and Balsam Poplar {Populus tremuloides, P. 
balsamifera). White Birch (Betula papyrifera), and 
Balsam Fir (Abies balsamea). Jack Pine {Pinus 
banksiana) occurs widely on sandy, drier sites such 
as eskers, moraines and outwash plains. Clay areas 
were cleared for agriculture in the 1930s and 1940s; 
these are limited in extent and distribution around 
communities on or very near Highway 11, which 
bisects the study area from southeast to northwest 
(Figure 1). Wetlands vary widely in size, and beaver 
impoundments on the many low gradient streams 
account for many of them. A wetland classification 
system for the area and an assessment of its rele- 
vance to duck species in the Clay Belt was devel- 
oped by Rempel et al. (1997) as part of this study. 

Population Estimation 

Distribution and abundance of breeding waterfowl 
in the Clay Belt was determined by aerial surveys of 
fixed plots during nest initiation periods in 1988, 
1989 and 1990. These plots were selected using a 
two-staged process in which the whole Clay Belt 
was divided into a grid of 10 X 10 km blocks follow- 
ing the UTM system. Each block was then subdivid- 
ed into 25 - 2 X 2 km plots, and one plot selected at 
random from each block for a total of 1 17 plots. This 
sampling regime allowed us to describe species dis- 
tribution, and to statistically estimate population size 
and habitat correlations. 

Surveys were carried out by helicopter (after Ross 
1985, 1987). All surveys were done from a Bell 
206B helicopter on high skids and equipped with a 
range extender on the fuel tank and bubble observa- 
tion windows on the back doors for improved visibil- 
ity. Two observers in the back watched for water- 
fowl from their respective sides of the aircraft, and 
passed observations by intercom to the navigator/ 
recorder in front. That person wrote these directly 
onto acetate-covered aerial photographs, recording 
species, sex, numbers, and exact locations of all 
waterfowl; care was taken to ensure that sightings 
came from within the boundaries of the plots. The 
aircraft passed over all suitable habitat in each plot at 
altitudes as low as 20 m, and at speeds ranging from 



a hover to lOOkm/h. Multiple passes were made 
over those waterbodies where the initial coverage 
was thought to be inadequate or where birds seen 
were not initially identified. Because most wetlands 
in the Clay Belt are small and clearly bounded, total 
coverage of all wetlands in the plots was accom- 
plished by following shorelines, and no transects 
were employed. All plots were surveyed during the 
following periods: 10-21 May 1988, 20-26 May 
1989, and 20-22 May 1990. 

Results of the survey are expressed as numbers of 
indicated breeding pairs (IBP) per species. For clear- 
ly dimorphic species, IBP was based on the total of 
lone males, pairs, and number of males in flocks 
with up to 5 males (as in Dzubin 1969). For Amer- 
ican Black Ducks {Anas rubripes) which are more 
weakly dimorphic, determination of sex was suc- 
cessful in about 90% of the cases; the remaining 
Black Ducks were ascribed indicated pair values 
based on the average proportions of indicated pairs 
to number of individuals in the three sighting classes 
(lone birds, two-bird groups, flock of three or more) 
as taken from all breeding pair surveys in northeast- 
ern Ontario from 1988 to 1990 (Ross, unpublished 
data). Indicated pairs for Canada Geese (Branta 
canadensis) were counted for each single, pair, or 
group of three. Counts of indicated pairs on the plots 
based on this helicopter survey technique are compa- 
rable to intensive ground survey results and are con- 
sidered sufficiently close to the real values to be 
used directly in breeding density calculations (Ross 
1985); no visibility correction factors have been 
applied. 

Habitat Parameters 

Although the Clay Belt is designated as a single 
forest section by Rowe (1972), other classifications 
can be used to subdivide this area to examine broad 
patterns in waterfowl distribution. In Figure 1, we 
present two classifications of the region. The first is 
an ecological classification showing the boundaries 
of the ecoregions and ecodistricts (Wicken 1986; 
Wickware and Rubec 1989); an ecodistrict is defined 
as a part of an eco-region characterized by a distinc- 
tive pattern of relief, geology, geomorphology, vege- 
tation, soils, water, and fauna. A second classifica- 
tion portrays substrate and is based on the surficial 
geology of the region (Glacial Map of Canada, Geo- 
logical Survey of Canada). To aid in interpretation of 
waterfowl distribution data, amounts of various 
potential nesting habitats (wetland associated with 
standing water) in each plot were measured by elec- 
tronic planimeter, and number of water bodies were 
counted. 

Statistical Analysis 

Overall breeding densities in both ecodistrict and 
substrate units were compared using a Mann- 
Whitney U Test (from Siegel 1956). Contour maps 



2002 Ross, Abraham, Gadawski, Rempel, Gabor, and Maher: Breeding Waterfowl 



45 



Table 1. Breeding densities of waterfowl of the Clay Belt (present and earlier studies) plus estimates of population sizes 
from present study. 





Dennis 


Ross 












Source 


(1974) 


(1987) 






Preseni 


t Study 












Mean 
















(1988- 




Year 


1973 


1982 


1988 


1989 


1990 


1990) 




Parameters 




Breeding Density (IBP/100 km^) 


Population Size (range) 


Species 


Canada Goose 


- 


1.0 


1.98 


3.24 


2.36 


2.52 


1330(1040-1710) 


Wood Duck 


- 


1.0 


0.54 


2.88 


2.17 


1.86 


980 (290-1520) 


Green-winged Teal 


9.7 


4.0 


4.50 


8.27 


11.96 


8.21 


4330(2370-6310) 


American Black Duck 


31.3 


10.2 


16.09 


19.85 


20.09 


18.72 


9870 (8490-10600) 


Mallard 


23.2 


21.0 


21.22 


29.50 


24.09 


24.88 


13120(11190-15560) 


Blue-winged Teal 


6.2 


6.0 


3.96 


7.01 


6.70 


5.88 


3100(2090-3700) 


American Wigeon 


17.8 


- 


2.16 


2.88 


3.99 


3.00 


1580(1140-2100) 


Other Dabblers 


- 


- 


- 


0.54 


0.18 


0.24 


130 (0-290) 


Ring-necked Duck 


12.4 


18.0 


16.01 


22.66 


24.09 


20.90 


11020(8440-12710) 


Lesser Scaup 


- 


- 


0.72 


0.72 


2.90 


1.44 


760 (380-1530) 


Common Goldeneye 


27.8 


11.0 


11.51 


11.70 


13.22 


12.11 


6390 (6070-6970) 


Bufflehead 


- 


- 


0.90 


0.90 


0.72 


0.84 


440 (380-470) 


Hooded Merganser 


3.1 


4.0 


6.65 


5.40 


8.15 


6.71 


3540 (2850-4300) 


Common Merganser 


- 


6.0 


5.58 


3.78 


5.25 


4.95 


2610(1990-2940) 


Other Divers 


1.3 


- 


- 


0.54 


0.18 


0.24 


130 (0-290) 


Total 132.8 82.2 


91.81 


119.8! 


) 126.07 


112.50 


59330 (48420-66490) 



of breeding pair densities and other parameters were 
generated using surfaces created by the potential 
mapping routine (POTMAP) of the SPANS geo- 
graphical information system (INTERA TYDAC 
Inc. 1991) and are based on sequential averaging of 
point values encompassed within an outer circle of 
47 km of the each specific plot value; an inner circle 
of 10 km holds the specific plot value constant over 
that area. 

Results and Discussion 

Waterfowl Population Sizes 

Breeding densities of the more common waterfowl 
species for each of the three years are presented in 
Table 1. Densities, for each species, were similar in 
1989 and 1990, but those for 1988 were consistently 
lower for dabbling ducks and Ring-necked Ducks 
(Aythya collahs). These differences reflected outlier 
values generated when very low counts were made 
in 1988 on a small number of plots that had much 
higher numbers in later years. Because annual varia- 
tion in waterfowl numbers in boreal systems is not 
well known and the ranges of values were less than 
20 percent of the average for the most common 
species, we used three-year means to represent aver- 
age breeding densities of waterfowl species in the 
Clay Belt (Table 1). 

Overall breeding density of waterfowl was 
1 12.5 IBP/ 100 km-; the four most common species 
(68% of total IBP) were American Black Duck, 
Mallard {Anas platyrhynchos). Ring-necked Duck, 



and Common Goldeneye {Bucephala clangula). Less 
common, but widely encountered (4 species, 23% of 
total IBP), were Green-winged Teal {Anas crecca). 
Blue-winged Teal {Anas discors). Hooded Mer- 
ganser {Lophodytes cucullatus), and Common 
Merganser {Mergus merganser). The remaining nine 
percent included five uncommon and locally occur- 
ring species [Canada Goose, Wood Duck {Aix spon- 
sa), American Wigeon {Anas americana). Lesser 
Scaup {Aythya ajfinis), Bufflehead {Bucephala albe- 
ola)], plus four rare species [Northern Pintail {Anas 
acuta). Northern Shoveler {Anas clypeata). Surf 
Scoter {Melanitta perspicillata). and White-winged 
Scoter {Melanitta fusca)]. Estimates of breeding 
populations of all but the rare species (also in Table 
1) were generated by multiplying average densities 
by the area of the Ontario Clay Belt (52 740 km-), to 
give an average total of 59 330 breeding pairs of 
waterfowl in this area (range: 48 420-66 490). 

Habitat Characteristics 

Only the two largest ecodistricts (Numbers 29 and 
30) had sufficient numbers of survey plots (81 and 
20. respectively; 86% of all plots) for separate analy- 
sis. Ecodistrict 30 had a lower elevation and a 
greater proportion of standing water than did 
Ecodistrict 29 (Wickware and Rubec 1989). Amount 
of waterfowl breeding habitat and number of water 
bodies per plot were significantly higher in Eco- 
district 30 (Table 2). Ecodistrict 29 contained most 
of the agricultural areas and much oi the area thai 
has been logged (Button and Black 1975). 



46 



The Canadian Field-Naturalist 



Vol. 116 



Surficial geology is another way of subdividing 
the Clay Belt (Figure 1). Although lacustrine sedi- 
ments, essentially clay, are the dominant substrate, 
much of the study area, particularly toward the west, 
is underlain by ground moraines with several smaller 
sections of end moraines. As clay usually contributes 
to greater ecological productivity, we expected that 
its presence should influence the distribution of 
some species. No statistical difference was found in 
number of water bodies or the total area of wetlands 
in plots of the two substrates (Table 2). 

Waterfowl Distribution 

Distributions of individual species are displayed as 
contoured maps based on mean breeding density per 
plot (Figure 2) for all species that were recorded on 
more than nine different plots. In Table 3, breeding 
densities of these species are compared between the 
two ecodistricts and between the two substrate types. 

The most abundant species was the Mallard that 
was found throughout the area with highest densities 
following a diagonal band (northwest-southeast) 
across the Clay Belt. There was no difference in 
breeding density between ecodistricts, but density 
was higher on plots with a clay substrate. Although 
habitat fertility influences Mallard distribution 
(Merendino and Ankney 1994), the area of highest 
breeding density is also that with most agricultural 
and tree-harvesting activities, which produce prairie- 
and parkland-like habitats to which Mallards are 
well-adapted (Bellrose 1980). 

American Black Ducks were abundant and 
widespread in the Clay Belt (second amongst dab- 
bling ducks, third overall). Highest breeding densities 
occurred in the north. Of the two ecodistricts, signifi- 
cantly higher breeding density was encountered in 
Ecodistrict 30, but unlike Mallards, no relationship 
was found between breeding density and substrate. 
Although Mallards and Black Duck are closely relat- 
ed and interbreed, their distributions did not show the 
same patterns even though they were among the most 
common species (see also Ross and Fillman 1990). 



The most common diving ducks in the Clay Belt 
(and second most abundant duck overall) were Ring- 
necked Ducks. Their density distribution was similar 
to that of Black Ducks, with which they regularly co- 
occur (McNicol et al. 1987). Ecodistrict 30 support- 
ed higher breeding densities, but there was no rela- 
tionship with substrate. 

Common Goldeneye was the fourth most abundant 
species, and like Black Ducks and Ring-necked 
Ducks, they occurred in significantly higher density 
in Ecodistrict 30 than in Ecodistrict 29, and there was 
no correlation between distribution and substrate. 

Green-winged Teal were observed throughout the 
study area but did not show a pattern explainable by 
either ecodistrict or substrate classifications, suggest- 
ing that the birds were using specific habitat types 
with distributions unrelated to these factors. In the 
Clay Belt, occurrence of this species is significantly 
associated with Beaver pond systems (Rempel et al. 
1997). Paquette and Ankney (1996) demonstrated 
high specificity in wetland use by this species in 
British Columbia. 

Hooded Mergansers occurred in moderate num- 
bers. High density zones largely coincided with 
those of Conmion Goldeneye, which may reflect the 
distribution of suitable nesting cavities for which the 
two species have similar requirements (Lumsden et 
al. 1980). No relationship with ecodistrict was evi- 
dent, but breeding density was significantly higher in 
areas with moraine substrate than in those underlain 
by clay; the latter substrate may produce greater tur- 
bidity in waterbodies making them less suitable for 
foraging by this species (Bellrose 1980). 

Blue-winged Teal distribution showed the same 
diagonal band component as that of Mallard with 
which it shares the ability to exploit more open habi- 
tats (Bellrose 1980). Numbers were too low, howev- 
er, to determine any relationship with either ecodis- 
trict or substrate type. 

Common Mergansers showed concentrations in 
those parts of the northern part of the Clay Belt that 



Table 2. Comparison of summary measures of waterfowl community parameters and habitat by ecodistrict and substrate. 



Classification 



Ecodistrict 



Substrate 



Significance Clay Moraine Significance 

(a) Waterfowl Community Parameters 29(n = 81) 30 (n = 20) (p) (n = 72) (n = 43) (p) 



Total Breeding Density 
(IP/100km2) 

Mean Number of 
Species per Plot 



95.08 



3.85 



151.14 



5.60 



NS 
(0.094) 

0.025 



115.00 



4.36 



97.46 



4.21 



NS 



NS 



(b) Habitat 


Mean Area per Plot of All 
Wetland Habitat (Ha) 

Number of Waterbodies per Plot 


20.75 
3.11 


27.97 
4.45 


0.003 
0.012 


21.53 
3.68 


21.91 

3.21 


NS 

NS 



2002 Ross, Abraham, Gadawski, Rempel, Gabor, and Maher: Breeding Waterfowl 



47 



Canada Goose 




IP/100 km 

□ 0-5 

□ 5-10 
M 10-20 
■ 20-40 



Mallard 



IP/100 tan 




n 



0-5 

5-10 

10-20 

20-40 

40-80 



American Black Duck 



IP/100 tan 




Green-winged Teal 



IP/100 tan 




Blue-winged Teal 




IP/100 ton 

□ 0-5 

□ 5-10 

B 10-20 
■ 20-40 



American Wigeon 




IP/100 ton 



D 
D 



0-5 

5-10 

10-20 

20-40 



Ring-necked Duck 



Common Goldeneye 



IP/100 Ion 





IP/100 ton 



□ 0-5 

□ 5-10 

■ 10-20 
H 20-40 

■ 40-80 



Hooded Merganser 




Number of Waterfowl 
Species per Plot 



IP/100 ton 



10-20 



20-40 
40-80 



Sp o d — Count 




Common Merganser 




Total Number of Waterfowl 




IP/100 ton 

□ 0-5 

□ 5-10 
■ 10-20 



IP/100 ton 



D 



25-50 

50-100 

100-200 

200-400 



Figure 2. Contoured maps showing breeding densities of individual waterfowl species, overall breeding density, and 
species richness 



48 



The Canadian Field-Naturalist 



Vol. 116 



Table 3. Comparisons between ecodistricts and between substrates of breeding densities of the major waterfowl species in 
the Clay Belt. 



Classification 




Ecodistrict 






Substrate 










Significance 






Significance 


Division 


No. 29 


No. 30 


(P) 


Clay 


Moraine 


(P) 


Species 


Mallard 


25.51 


20.00 


NS 


29.17 


14.92 


0.013 


Ring-necked Duck 


12.35 


37.92 


< 0.001 


20.95 


17.05 


NS 


American Black Duck 


13.81 


29.47 


0.008 


16.39 


18.00 


NS 


Common Goldeneye 


8.85 


21.25 


< 0.001 


12.38 


10.85 


NS 


Green- winged Teal 


8.54 


7.50 


NS 


8.45 


9.30 


NS 


Hooded Merganser 


6.69 


6.67 


NS 


5.09 


9.88 


0.04 


Blue-winged Teal 


5.45 


6.25 


NS 


6.48 


3.29 


NS 


Common Merganser 


2.88 


6.67 


NS 


3.82 


5.04 


NS 


Canada Goose 


2.06 


8.33 


0.006 


2.89 


3.29 


NS 


American Wigeon 


3.70 


1.25 


NS 


3.70 


1.55 


NS 



had high concentrations of Black Ducks and Ring- 
necked Ducks. No relationship with either substrate 
or ecodistrict could be demonstrated, probably due to 
the small numbers recorded. Common Mergansers 
have shown an affinity for rivers (McNicol et al. 
1987; Rempel et al. 1997) in the Boreal Forest and 
their distribution in the Clay Belt may reflect the 
concentration of tributaries that converge toward the 
Moose River in the northeast. 

Small numbers of Canada Geese were found north 
of Lake Abitibi and Highway 1 1 in habitat showing 
strong muskeg elements of the Hudson Bay Low- 
lands which lie immediately to the north, and are the 
centre of the breeding range of the Southern James 
Bay Population of Canada Geese (Raveling and 
Lumsden 1977). Breeding density was significantly 
higher in Ecodistrict 30. 

American Wigeon were uncommon and not 
recorded in sufficient numbers to define distribution 
or determine regional differences. The species 
prefers larger permanent marshes with much open 
water (Palmer 1976) and shows an affinity for lake 
estuary marsh in the Clay Belt (Rempel et al. 1997). 
Northern river deltas held the highest breeding den- 
sities of this species (Bellrose 1980). 

Overall Waterfowl Distribution and 
Species Richness 

Contoured maps of total breeding pair density of 
waterfowl and numbers of waterfowl species per plot 
(Figure 2) indicated that both overall waterfowl den- 
sity and species richness were higher in the north. 
However, there was a significant relationship only 
between ecodistrict and species numbers (higher in 
Ecodistrict 30; Table 2). This may be an "edge 
effect" (i.e., higher habitat diversity) due to the inter- 
gradation with the Hudson Bay Lowlands immedi- 
ately to the north. Another area of higher species 
richness and density occurred around Timmins and 
Lake Abitibi near the south-eastern edge of the Clay 



Belt, possibly reflecting greater habitat diversity 
where the Clay Belt grades into the Great Lakes-St. 
Lawrence Forest Region. Lowest species richness 
and density occurred around the southwestern edge 
in a very sandy section of the Chapleau Plain. 

Comparison with Other Studies 

Our survey showed that waterfowl breeding in the 
Clay Belt were not distributed uniformly. Species 
richness was significantly higher towards the north, 
particularly in Ecodistrict 30, and total numbers of 
breeding pairs showed a similar tendency. This gra- 
dient towards the north was significant for four 
species (Canada Goose, Black Duck, Ring-necked 
Duck, and Common Goldeneye), all of which are 
characteristic of boreal habitats and seem to respond 
to physiography, particularly the amount of wetland 
habitat. Mallards and Blue-winged Teals were con- 
centrated in a northwest-southeast strip bisecting the 
Clay Belt which correlates with the distribution of 
clay substrate. These species are more broadly dis- 
tributed in other habitats to the south and west and 
may be responding either directly or indirectly to the 
higher productivity associated with clay (Merendino 
and Ankney 1994). Other species, such as the 
American Wigeon and the Green-winged Teal, did 
not exhibit either of these patterns and instead may 
be responding to distributions of very specific habi- 
tat types. 

Initial comparison of our results with those of ear- 
lier surveys (Table 1) indicated substantial differ- 
ences. However, the previous surveys only covered 
part of the Clay Belt and locations of survey plots in 
earlier studies strongly influence the results. The 
Clay Belt plots used by Dennis (1974) were mostly 
in the relatively wetland-rich Ecodistrict 30 and 
other areas north of Highway 11, where one would 
expect a higher total breeding density than the aver- 
age from the present survey. Conversely, the survey 
by Ross (1987), that yielded a relatively low total 






2002 Ross, Abraham, Gadawski, Rempel, Gabor, and Maher: Breeding Waterfowl 



49 



breeding density, was restricted to a 100 X 100 km 
block in the south central portion of the Clay Belt 
and included areas with the lowest breeding densities 
in this study. These relationships hold for three of 
the most common species. Black Duck and Common 
Goldeneye densities were highest in the Dennis 
(1974) survey while Black Duck density was lowest 
in the Ross (1987) count. Both species demonstrate 
the previously noted north-south density gradient. 
Densities for the Mallard, which did not show the 
north-south gradient, differed little among surveys. 
This implies that there have not been major changes 
in the Clay Belt populations of these three species 
since 1973. Of the more abundant species, only the 
Ring-necked Duck showed results [lowest count in 
the Dennis (1974) survey] that are not explainable by 
their present distribution. 

Total waterfowl breeding density is slightly higher 
on the Clay Belt than on the surrounding boreal forest 
characterized by exposed Precambrian Shield 
although the ranges of values partially overlap [Clay 
Belt (Table 1): 91.8-126.1 IBP/100 km2 vs. Exposed 
Shield (Ross 1987): 75.0-107.8 IBP/100 km2]. 
Species composition, however, differs considerably. 
Generally, dabblers such as the Mallard, and Blue- 
winged Teal are more common in the Clay Belt 
whereas the mergansers attain higher densities on the 
exposed shield; goose numbers are minimal in both 
areas. The higher density of wetlands in the exposed 
shield appears to compensate in part for the higher 
productivity of individual wetlands in the Clay Belt 
(Ross 1987). The Hudson Bay Lowland immediately 
to the north supports, lower densities of ducks 
(approx. 37 IBP/100 km^; Ross, unpubhshed informa- 
tion) but higher numbers of geese (5-13 IBP/100 km^ 
for Canada Geese). Overall the contribution of the 
Clay Belt to northern Ontario duck numbers is essen- 
tially proportional to its area. 

Conservation Implications 

Better understanding of distribution of waterfowl 
and characteristics of their breeding habitat allows 
more focussed and effective conservation actions. 
Areas being considered for acquisition and protection 
can be priorized based on waterfowl breeding density 
and species richness. Numbers of waterfowl potential- 
ly at risk due to large-scale developments, (e.g., 
forestry, hydro-electric dams, and mining) can be 
determined and remedial action plans prepared. 
Furthermore, some insight can be gained into the 
effects of specific management actions. For example, 
efforts to increase the amount of wetland habitat may 
well show colonization by species such as Black Duck, 
Ring-necked Duck, Common Goldeneye, and possibly 
Canada Goose. If this managed habitat also has a rich 
clay substrate, one can also expect an increase in 
Mallards (and possibly Blue-winged Teal); however, 
this may lead to competitive interaction between Black 
Ducks and Mallards (Merendino and Ankney 1994). 



Target species and substrate type should be carefully 
considered in determining site locations for habitat 
management activities in the Clay Belt. 

Acknowledgments 

Financial support was provided by Ducks 
Unlimited Canada, the Ontario Ministry of Natural 
Resources through the Northern Forest Development 
Group, Wildlife Habitat Canada, and the Canadian 
Wildlife Service. We are very grateful for the field 
assistance provided by M. Barr, P. Burke, B. Di 
Labio, D. Fillman, D. Kopenhaver, D. Morgan, S. 
and S. O'Donnell, M. Porter, W. Simkin, N. Wilson, 
and J. Woodcock. J. Boos, R. Clay, and D. Kroeker 
of Ducks Unlimited staff at various times all made 
important contributions. We thank R. Watt of the 
OMNR for his support of this study and also the 
OMNR field offices whose radio operators provided 
flight following for our aircraft. We greatly appreci- 
ate the work of B. Campbell in data processing and 
figure preparation. Helpful comments were received 
from A. J. Erskine and an anonymous reviewer. 

Literature Cited 

Addy, C. E., W. F. Crissey, H. R. Webster, and G. F. 
Boyer. 1952. Waterfowl breeding ground survey in east- 
ern Canada. U.S. Fish and Wildlife Service Scientific 
Report Number 21.81 pages. 

Bellrose, F. C. 1980. Ducks, Geese and Swans of North 
America. 3rd edition. Stackpole Books, Harrisburg, 
Pennsylvania. 540 pages. 

Chamberlain, E. B., and C. F. Kaczynski. 1965. Prob- 
lem in aerial surveys of waterfowl in eastern Canada. 
U.S. Fish and Wildlife Service Special Scientific Report 
— Wildlife Number 93. 21 pages. 

Dennis, D. G. 1974. Waterfowl observations during the 
nesting season in Precambrian and Clay Belt areas of 
north-central Ontario. Pages 53-96 in Canadian Wildlife 
Service Waterfowl Studies in Eastern Canada 1969-73. 
Edited by H. Boyd. Canadian Wildlife Service Report 
Series Number 29. 

Dzubin, A. 1969. Assessing breeding populations of 
ducks by ground counts. Pages 178-230 in Saskatoon 
Wetland Seminar. Canadian Wildlife Service Report 
Series Number 6. 

Hanson, H. C, M. Rogers, and E. S. Rogers. 1949. 
Waterfowl of the forested portions of the Canadian 
Precambrian Shield and the Palaeozoic Basin. Canadian 
Field-Naturalist 63: 183-204. 

Hutton, C.L. A., and W. A. Black. 1975. Ontario Arctic 
Watershed. Lands Directorate Map Folio Number 2. 
Environment Canada. 107 pages. 

Kaczynski, C. F., and E. B. Chamberlain. 1968. Aerial 
surveys of Canada Geese and Black Ducks in eastern 
Canada. U.S. Fish and Wildlife Service Special 
Scientific Report Number 188. 29 pages. 

Lumsden, H. G., R. E. Page, and M. Gauthier. 1980 
Choice of nest boxes by Common Goldeneyes in 
Ontario. Wikson Bulletin 92: 497-505. 

McNicoI, D. K., B. E. Bendell, and R. K. Ross. 1987. 
Studies of the effects of acidification on aquatic wildlife 
in Canada: waterfowl and trophic relationships in small 
lakes in northern Ontario. Canadian Wildlife Service 
Occasional Paper Number 62. 76 pages. 



50 



The Canadian Field-Naturalist 



Vol. 116 



Merendino, M. T., and C. D. Ankney. 1994. Habitat use 
by Mallards and American Black Ducks in central 
Ontario. Condor 96: 41 1-421. 

Palmer, R. S. Editor. 1976. Handbook of North American 
Birds, Volume 2. Yale University Press, New Haven, 
Connecticut. 521 pages. 

Paquette, G. A. and C. D. Ankney. 1996. Wetland selec- 
tion by American Green-winged Teal breeding in British 
Columbia. Condor 98:27-33. 

Raveling, D. G. and H. G. Lumsden. 1977. Nesting ecolo- 
gy of Canada Geese in the Hudson Bay Lowlands of 
Ontario: evolution and population regulation. Ontario 
Ministry of Natural Resources Fish and Wildlife 
Research Report Number 98. 77 pages. 

Rempel, R. S., K. F. Abraham, T. R. Gadawski, T. S. 
Gabor, and R. K. Ross. 1997. A simple wetland classi- 
fication for boreal forest waterfowl. Journal of Wildlife 
Management 61: 746-757. 

Ross, R. K. 1985. Helicopter vs. ground surveys of water- 
fowl in the boreal forest. Wildlife Society Bulletin 13: 
153-157. 

Ross, R. K. 1987. Interim report on waterfowl breeding 
pair surveys in northern Ontario, 1980-83. Canadian 
Wildlife Service Progress Note 168. 8 pages. 



Ross, R. K., and D. Fillman. 1990. Distribution of 

American Black Duck and Mallard in northern Ontario. 

Canadian Wildlife Service Progress Note 189. 5 pages. 
Rowe, J. S. 1972. Forest Regions of Canada. Canadian 

Forestry Service Publication 1300. 172 pages. 
Sharpe, J. F., and J. A. Brodie. 1931. The Forest 

Resources of Ontario. Department of Lands and Forests, 

Toronto. 
Siegel, S. 1956. Nonparametric Statistics for the Behavioral 

Sciences. McGraw-Hill Book Company, New York. 312 

pages. 
Wellein, E. G., and H. G. Lumsden. 1964. Northern forest 

and tundra. Pages 67-76 in Waterfowl Tomorrow. 

Edited by J. P. Linduska. U.S. Department of the 

Interior, Washington, D.C. 
Wicken, E. 1986. Terrestrial ecozones of Canada. Eco- 
logical Land Classification Series 19. Environment 

Canada, Ottawa. 26 pages. 
Wick ware, G. M., and C. D. A. Rubec. 1989. Ecoregions 

of Ontario. Ecological Land Classification Series 

Number 26. Environment Canada, Ottawa. 37 pages. 

Received 12 October 2000 
Accepted 3 May 2002 



Resilience of Foothills Rough Fescue, Festuca campestris, 
Rangeland to Wildfire 

Edward W. Borki, Barry W. Adams^, and Walter D. Willms^ 

1 Range Scientist, Department of Agricultural, Food, and Nutritional Science, University of Alberta, 410E, AgFor Centre, 

Edmonton, Alberta T6G 2P5 Canada, 
2Range Management Specialist, Public Lands Division, Alberta Agriculture, Food, & Rural Development, Bag 3014, 

Agriculture Centre, Gaol Road, Lethbridge, Alberta TIJ 4CJ Canada 
3Range Ecologist, Lethbridge Research Centre, Agriculture and Agri-Food Canada, P.O. Box 3000, Lethbridge, Alberta 

TIJ 4B1 Canada 

Bork, Edward W., Barry W. Adams, and Walter D. Willms. 2002. Resilience of Foothills Rough Fescue, Festuca 
campestris, rangeland to wildfire. Canadian Field Naturalist 116(1): 51-59. 

A three year monitoring program evaluated the effects of a December 1997 wildfire in southwest Alberta, on Foothills 
Rough Fescue grassland species composition, ground cover, herbage production, and forage quality. Changes in species 
abundance included a reduction in grass cover (p<0.10) after burning. Rough Fescue also increased seedhead production 
during the second year after fire (p = 0.08). Relative to the unbumed area, graminoid production declined (p<0.05) by 
approximately 40% with burning, while forb production was unaffected. By the second growing season, live plant cover 
and herbage production had recovered on the burned area. The forage quality of individual Foothills Rough Fescue plants 
was greater on the burned area, with the greatest increase in crude protein in 1998 (p<0.10), and energy and total 
digestibility in 1999 (p < 0.05). Increased quality may be linked to the level of forage production, as well as a fire-induced 
delay in plant phenology. Although soil erosion appeared to be minimal, there was an increase in exposed soil and a corre- 
sponding decline in litter and mulch cover (p< 0.05). Greater nitrogen levels (p = 0.051) were found in creeks downstream 
of the bum area during 1998, indicating some nutrient losses may be attributed to the fire. Although the grasslands exam- 
ined displayed considerable resilience to this severe wildfire, favourable recovery was probably linked to the high precipi- 
tation during 1998, when summer rainfall was 48% above average. 

Key Words: Foothills Rough Fescue, Festuca campestris, forage quality, precipitation, production, resilience. Alberta 



Foothills Rough Fescue {Festuca campestris 
Rydb.) grasslands are well-adapted to fire, having a 
historical fire return interval of 5-10 years (Wright 
and Bailey 1982). During the last century, however, 
fragmentation of rangeland landscapes and fire sup- 
pression is thought to have enhanced the risk of 
severe fires by lengthening fire return intervals and 
increasing fuel accumulation during the burn inter- 
cession. Antos et al. (1983) concluded that Foothills 
Rough Fescue grasslands were best adapted to mod- 
erate frequency fire (e.g., every 6 years). 

From January of 1997 through August of 2000, 
eight known wildfires were documented in the 
Fescue Prairie region of Alberta affecting nearly 
33 500 ha, with at least five of these greater than 400 
ha (Barry Adams, personal observation). This trend 
prompted concerns over the ability of these grass- 
lands to recover from wildfire, particularly those fol- 
lowing the prolonged absence of fire. Native rough 
fescue grasslands may be less tolerant of infrequent 
fire due to the increased severity of burning associat- 
ed with litter accumulation. Rough fescue in particu- 
lar, is of value to the ranches in the region because of 
its role in providing a practical and economical 
source of fall and winter grazing (Willms et al. 
1993), as well as habitat for wildlife. 

Although several studies investigating the response 



of fescue grassland to fire have been conducted, these 
typically involve Plains Rough Fescue {Festuca hallii 
(Vasey) Piper) grasslands in the Aspen Parkland of 
Alberta and Saskatchewan (Bailey and Anderson 
1978; Anderson and Bailey 1980; Redmann et al. 
1993; Gerling et al. 1995). Furthermore, these studies 
examine areas burned under prescription rather than 
wildfire. Antos et al. (1983) examined changes in 
species composition following a small (49 ha), uncon- 
trolled wildfire in July within a Foothills Rough 
Fescue-Idaho Fescue {Festuca idahoensis Elmer) 
plant community in Montana. 

As a result of concerns associated with increasing- 
ly common wildfire events in Alberta and the lack of 
information on their ecological impacts, a research 
plan was formulated to assess Foothills Rough 
Fescue grassland recovery following a large-scale 
wildfire that occurred in December 1997. Initial 
inspection of the burned area indicated that many 
rough fescue plants had been heavily damaged 
and/or killed, presenting a unique opportunity to 
examine the impact of a dormant season fire on the 
resilience of these grasslands. Specific objectives of 
this project were to determine the short-term (2-3 
year) impacts of the fire on: (1) plant community 
composition and ground cover, (2) above-ground net 
primary production, and (3) rough fescue forage 



51 



52 



The Canadian Field-Naturalist 



Vol. 116 



quality. A secondary objective was to investigate the 
area for soil erosion. 

Methods 

Study Area 

The study area was burned by a wildfire on 14 
December 1997, covering 220 km^ southwest of 
Granum, Alberta (113°50"W; 49°45"N) within the 
Fescue Prairie Ecoregion (Strong 1992). The fire 
began around 11:00 and traversed 33 km in under 
four hours. The fire was preceded by a dry autumn 
and aided by sustained winds of 30-40 km/hr, gust- 
ing to 70 km/hr. Weather conditions at the time of 
the fire were unusual for December, with a maxi- 
mum mid-day temperature of 13°C (10°C above 
average) and relative humidity of 17% (Tymstra 
1998*). Although detailed fuel-load data are 
unavailable for the burned area itself, litter loads 
adjacent to the burn averaged nearly 900 kg/ha. 
Approximately 83% of the 21 600 ha burn affected 
Foothills Rough Fescue grasslands. The economic 
and social impacts of this fire on the ranching com- 
munity within the region were widely publicized at 
the time. 

A fire intensity assessment prepared by Tymstra 
(1998*) described the Granum fire as extremely hot 
with a head fire intensity ranging from 10 000 to 
20 000kW/m2. Tymstra (1998*) further concluded 
that the average rate of fire spread (-10 km/hr) was 
one of the greatest documented for grassland fires in 
Canada. 

Topography of the area varied from steep foothills 
to gently sloping terraces, with occasional flat valley 
bottoms. Elevations range between 1 000 and 1 500 
m. Native grasslands are dominated by mid to late 
successional Foothills Rough Fescue-Parry Oatgrass 
(Danthonia parryi Scribn.) communities, interspersed 
with riparian vegetation along wetlands and streams, 
and xeric grasslands dominated by Idaho fescue and 
Parry Oatgrass on sites with shallow or poorly devel- 
oped soils. In some areas, livestock grazing has 
resulted in an increase in introduced species such as 
Kentucky Bluegrass {Poa pratensis L.) (Willms et al. 
1995, 1996). The dominant soil type is an Orthic 
Black Chernozem (Johnston et al. 1971). 

Field Procedures 

In May 1998, 10 sites were selected within 
Foothills Rough Fescue-Parry Oatgrass (Festuca- 
Danthonia) communities for monitoring, with eight 
situated along the perimeter burn as paired burned- 
unbumed sampling sites, increasing the likelihood of 
evaluating vegetation responses to the fire itself. All 
paired sites were located on a uniform ecosite (i.e., 
aspect and soils) and were intersected by a human- 
made (i.e., grader-bladed) fire boundary. This strati- 
fication helped ensure that vegetational differences 
between paired transects were caused by the fire 



rather than a naturally occurring difference in physi- 
cal site characteristics. Two additional sites were 
sampled within the interior of the bum. 

At each site, a 30-m linear transect was randomly 
established and permanently marked to facilitate re- 
sampling in later years. On each transect, ocular esti- 
mates of canopy cover were done on all plant species 
within 15 systematically placed 0.1 -m^ quadrats 
(Daubenmire 1959) at peak vegetative growth in June 
of each year. In addition, the cover of loose litter and 
mulch, as well as the amount of bare soil were esti- 
mated. Mulch was defined as the matted layer of fine 
organic material overlying mineral soil, consisting of 
heavily degraded litter in combination with fine roots 
from plants, either dead or alive (Dix 1960). 

Herbage production was determined within four, 
1.5 m by 1.5 m portable cages randomly set up along 
each transect. During the last week of August in each 
year, all current above-ground net primary produc- 
tion (ANPP) was clipped within a 0.5-m2 quadrat in 
each cage. Cages were moved to new locations 
between years. All ANPP was sorted to graminoid, 
forb, and litter components, dried at 50°C for 3 days, 
and weighed to determine dry matter. Rough fescue 
seedhead counts were done the first week of August 
within 15, l-m^ quadrats nested over the Dauben- 
mire quadrats along each transect to assess the 
potential for seed production in each year. 

To evaluate the effect of burning on forage quali- 
ty, individual rough fescue plants were randomly 
selected and harvested during the first week of 
August. In 1998 and 1999, five and eight plants, 
respectively, were clipped on each transect. After 
standing dead material was removed, plant samples 
were analysed for crude protein (CP), acid detergent 
fiber (ADF), and total digestible nutrients (TDN), as 
these variables are important for evaluating the qual- 
ity of forage available to livestock and wildlife 
(Holechek et al. 1998). In the third growing season 
after burning, vegetation sampling was limited to 
seedhead counts, the evaluation of exposed bare soil, 
determination of ANPP, and litter mass. 

Watershed-level impacts of the fire were assessed 
from water samples taken within three creeks that 
flowed through the burn area. Samples were taken 
immediately above and below the bum between 22 
January and 23 March, 1998 during snowmelt. All 
samples were analysed for total suspended solids 
(TSS), total dissolved sediment (TDS), and nitrogen 
(N) (Eaton et al. 1995). 

Data Analysis 

Direct comparisons were made using paired t-tests 
between the bumed and unbumed transects along the 
fire boundary on species richness (number of species), 
the cover of major grasses including foothills rough 
fescue and the wheatgrasses (Agropyron spp.), bare 
soil, litter, and mulch. The wheatgrasses [Northern 



2002 



BoRK, Adams, Willms: Resilience of Foothills Rough Fescue 



53 



(Agropyron dasystachyum (Hook.) Scribn.), Western 
{Agropyron smithii Rydb.), and Bearded {Agropyron 
subsecundum (Link) A.S. Hitchc.)] were pooled dur- 
ing sampling due to difficulty in identifying each 
species based on vegetative characteristics, particular- 
ly within the recovering bum area. Total legume cover 
was also assessed because these species constitute a 
key functional group responsible for N-fixation. 
Separate t-tests were done on the 1998 and 1999 data 
to evaluate the extent of recovery through time. 
Although data from the two transects in the interior of 
the bum could not be directly (i.e., statistically) com- 
pared with those from the bum perimeter, data from 
these sites were averaged for comparison to the other 
locations. Graminoid and forb ANPP, litter, and 
Rough Fescue seedhead counts, along with forage 
quality parameters, were also tested for burning 
effects in 1998, 1999, and 2000. 

Results and Discussion 

Growing Conditions 

Precipitation the year after the fire was favourable 
for recovery, with the majority falling during the 
growing season (Figure 1). Precipitation in 1998 at 
the nearby Agriculture Canada research station near 
Stavely was 502 mm from May to August, with a 
total of 647.5 mm in 1998, 46% above the regional 
average reported by Environment Canada (unpub- 
lished data). These data corroborate records from a 
ranch situated within the burn area, where April 
through August rainfall totalled 538 mm, and annual 
precipitation totalled 771 mm. In 1999, growing con- 



ditions approximated the norm for the region, with a 
total of 285 mm from May to August and annual pre- 
cipitation of 399 mm (Figure 1). 

Landscape patterns of green-up were variable in 
1998, with some areas developing rapidly, as evi- 
denced by advanced vegetative growth and the emer- 
gence of Rough Fescue seedheads by early May. 
Plant development appeared to be particularly 
advanced at the perimeter of the bum, possibly due 
to increased soil temperatures associated with the 
removal of litter (Antos et al. 1983), which could 
hasten tiller development. In other areas, however, 
phenology was noticeably delayed. Variation in veg- 
etation development is likely attributed to differ- 
ences in ecosite conditions such as slope, aspect, and 
soils, as well as corresponding variation in fire 
behaviour across the landscape. General reconnai- 
sance suggested that the slowest development 
occurred within the centre of the burn where the 
greatest evidence of fire intensity was observed both 
during the fire (i.e., in terms of flame length and rate 
of spread) and after the burn (i.e., surface distur- 
bance). At these locations, intense heat may have 
penetrated more deeply into the soil, increasing dam- 
age to plant meristems and carbohydrate reserves by 
destroying plant tissue. Greater damage, in turn, 
would force plants to resume growth from perennat- 
ing buds, slowing growth. In 1999, the phenological 
development of burned vegetation was uniform 
throughout the bumed area and more similar to that 
of the adjacent unbumed vegetation. 



iJan. -April 



May-June 



nJuly-Aug. 



n Sept. -Dec. 



400 

350 

^ 300 

^ 250 

c 

I 200 

% 150 

<D 

^ 100 

50 









kjtl] 




1997 



1998 



1999 



2000 



40 Year Ave 



Figure I . Yearly precipitation from January 1997 to December 2000 for the Agriculture Canada Stavely sub-station, locat- 
ed approximately 50 km north of the December 1997 wildfire, and comparison to long-term average precipitation 
from the Claresholm-Meadow Creek, Alberta climate station (Environment Canada, unpublished data). 



54 



The Canadian Feld-Naturalist 



Vol. 116 



Plant Species Abundance and Ground Cover 

A summary of the differences in plant species 
abundance in 1998 and 1999 among each of the 
three sample locations (unburned, perimeter burn, 
and interior bum) is provided in Table 1. Differences 
between burned and unburned sites along the fire 
boundary are consistent with the notion that fire is 
effective in changing plant community composition 
(Daubenmire 1968). 

Species richness (number/1.5 m^) and overall 
diversity were similar between burned and unburned 
areas (Table 1). Although the cover of introduced 
species tended to increase by the end of the second 
growing season (Table 1: 1999 data), this change was 
not significant (p>0.10). Concerns that Kentucky 
Bluegrass, due to its strongly creeping growth habit, 
might be more tolerant of fire and capable of invad- 
ing areas where other plants had been removed by 
fire, did not appear to be substantiated. At two sites, 
Kentucky Bluegrass, and to a lesser extent Dandelion 
(Taraxacum officinale Weber), did increase indicat- 
ing the expansion of invasive species may be linked 
to their presence and abundance within the communi- 
ty prior to burning. 

Only total grass cover was reduced within the 
burned area relative to the unburned in 1998 
(p<0.10; Table 1), with recovery by 1999. In com- 
parison to the perimeter bum, grass cover was less 
but forb cover greater, within the burn interior. 
Species data indicate the forb response was likely 
due to an increase in legumes such as Lupine 
(Lupinus argenteus Pursh), Locoweed (Oxytropis 
sericea Nutt.), Buffalo Bean (Thermopsis rhombifo- 
lia R.Br.), and Vetchling (Vicia americana Muhl), 
all of which were more abundant in both sampling 
years (Table 1). Anderson and Bailey (1980) also 
found increases in 4 of 5 legume species associated 
with annual prescribed burning in Plains Rough 
Fescue grasslands of the Parkland in central Alberta. 
Other research has indicated legumes are well adapt- 
ed to fire and increase following buming due to the 
termination of seed dormancy (Martin et al. 1975). 
Although not tested here directly, this factor may 
contribute to the observations made in this study and 
merit further investigation within Foothills Rough 
Fescue grasslands. Regardless of the mechanism, the 
increase in legumes is important in that they con- 
tribute nitrogen through symbiotic N-fixation, which 
may in turn, facilitate plant community recovery fol- 
lowing fire. 

Of the dominant grasses. Parry oatgrass appeared 
to decline the most the year after fire, followed by 
the wheatgrasses (Table 1). In contrast, sedges 
{Carex spp.) and Foothills Rough Fescue appeared to 
be resilient, with minimal difference in canopy cover 
across the fireline (Table 1). Rough fescue, however, 
had notably less cover at the interior of the bum rela- 
tive to the perimeter. Assuming these transects had 



similar amounts of Rough Fescue as the other areas 
sampled (this assumption is supported by the pres- 
ence of numerous dead fescue plants in the interior), 
buming may have been more severe at this location. 
The interior bum transects were generally located in 
areas where previous grazing may have been less 
intense, leading to greater litter accumulation. This 
interpretation is supported by the results of a 1990 
range survey that described the grazing history of the 
interior bum area as very light, and was mapped as 
secondary range with considerable litter build up 
(Tannas 1990). 

With the exception of Parry Oatgrass, all domi- 
nant grasses appeared to recover by the second year. 
The resilience of Foothills Rough Fescue following 
buming is inconsistent with other studies document- 
ing fire-induced declines in rough fescue (e.g., 
Bailey and Anderson 1978; Antos et al. 1983). Most 
previous studies, however, have examined Plains 
Rough Fescue rather than Foothills Rough Fescue, 
which differ in morphology (Pavlick and Looman 
1984), and presumably, their tolerance to fire. 
Mitchell (1957) suggested that the coarse stubble of 
rough fescue normally insulated perennating buds 
near the soil surface. With dry conditions, heavy 
stubble may allow the development of hot fires that 
burn into plant crowns as observed at the interior 
study locations. Antos et al. (1983) reported severe 
damage and high mortality of rough fescue on 
ungrazed sites where low fire frequencies allowed 
heavy litter accumulation, with three years needed 
for fescue recovery. 

In this investigation, the wheatgrasses declined 
more than Rough Fescue despite being generally 
considered better adapted to tolerate fire due to their 
rhizomatous growth habit (Wright and Bailey 1982). 
Other studies of wheatgrass response to burning 
show varied results, although most research has been 
done in drier regions. In some, fall buming increases 
wheatgrasses (Wright 1974; White and Currie 1983), 
while in others, wheatgrasses decline (Erichsen- 
Arychuk et al. 2002). The response observed here 
could also be due to the presence of Bearded 
Wheatgrass within the stand, which lacks rhizomes 
and may be susceptible to fire. 

Exposed Soil, Litter Cover, and Erosion Losses. 

Cover of bare soil, litter, and mulch varied with 
sampling location (Table 1). In general, litter was 
significantly (p<0.05) less across the fire-line on the 
bumed area from 1998 through 2000, with a corre- 
sponding increase in bare soil and mulch. By 2000, 
bare soil was no longer different between adjacent 
bumed (3.4%) and unbumed areas (1.4%), likely due 
to plant recovery and the re-accumulation of litter. 
Bare soil was significandy (p <0.05) less across the 
fire boundary in 1999 but not 1998 (p>0.10), and 
may indicate some drying and loss of protective 
mulch on the burned area between sampling periods. 



2002 



BoRK, Adams, Willms: RESiLffiNCE of Foothills Rough Fescue 



55 



Table 1 . Mean canopy cover (%) of major plant species and functional groups at each location sampled in 1998 and 1999. 
Species listed only include those with a minimum average cover of 1%. Values in parentheses represent one SE for vari- 
ables with significance tests. 





Canopy Cover— 1998 


Canopy Cover — 1999 


Species 


Interior Bum 


Perimeter Bum 


Unbumed 


Interior Bum 


Perimeter Bum 


Unbumed 


Agropyron spp. 


2.1 


6.5 


9.1 


12.4 


15.5 


11.9 


Carex spp. 


1.3 


7 


7.8 


4.2 


16.7 


13.3 


Danthonia parryi 


0.4 


0.8 


9.2 


2.5 


4.1 


13.7 


Festuca campestris 


3.6 


10.1 


11.6 


25.5 


22.4 


23.5 


Festuca idahoensis 




4.1 


5.7 


0.2 


13.9 


9.3 


Helictotrichon hookeri 




0.9 


0.8 




5.2 


5.1 


Koeleria macrantha 




0.8 


2.5 




3.3 


1.8 


Poa pratensis 


0.2 


0.3 


0.5 


3.3 


3.2 


0.6 


Poa sandbergii 










4.2 


2.4 


Stipa viridula 


0.2 






2.3 


4.8 




Total Grass: 


7.8 


30.5 a (3.8)* 


47.5 b (5.6) 


50.4 


93.4 a (4.8) 


82.0 a (7.8) 


FoRBs: 














Achillea millefolium 


1.2 






7.1 




0.1 


Agoseris glauca 


1 


0.9 


2.1 


4.4 


4.2 


3.9 


Androsace septentrionalis 








6.2 


0.6 




Anemone multifida 


1.1 


2.8 


L7 


3.1 


4 


2.2 


Aster laevis 


0.6 


0.3 


0.1 


3.1 


0.1 


0.6 


Astragalus pectinatus 




0.1 


0.5 


5.1 


0.9 


1.6 


Cerastium arvense 


0.1 


0.2 


0.8 


0.5 


1.4 


1.2 


Commandra pallida 




0.3 


0.7 




1.5 


1.6 


Companula rotundifolia 








2.9 


0.5 


0.5 


Erigeron caespitosum 




2.2 


1.2 




6.3 


2.1 


Galium boreale 


1.7 


2.6 


1.6 


5 


4.2 


4.7 


Geum triflorum 


1.6 


0.1 




2.2 


0.3 




Heterotheca villosa 




1 


1.3 


1 


3.4 


2.8 


Liatris punctata 




0.3 


0.8 




1.1 


1.8 


Lomatium dissecta 


0.2 


0.3 


0.3 


0.7 


0.3 


1.1 


Lupinus argenteus 


8.8 


L3 


1.8 


16.2 


2.7 


4.3 


Oxytropis sericea 




0.8 


0.7 




3.3 


1.4 


Phlox hoodii 




0.1 


0.7 




0.4 


1.2 


Solidago missouriensis 




0.1 




0.7 


2.8 


0.9 


Taraxacum officinale 


0.1 


0.3 


0.4 


0.4 


0.7 


0.3 


Thermopsis rhombifolia 


10.4 


2.8 


4.5 


21.4 


10.7 


9.3 


Tragopogon dubius 


0.1 


0.2 


0.1 


0.2 


0.1 


1.1 


Vicia americana 


0.6 


1.3 


0.4 


0.1 


2 


1.3 


Viola canadensis 


0.7 


0.1 




0.4 


1 




Zygadenus venenosus 


0.1 


0.9 


0.2 


0.6 


1.3 


0.1 


Total Forb: 


28.7 


20.6 a (4.5) 


22.2 a (3.6) 


82.1 


55.9 a (5.4) 


46.8 a (5.6) 


Legume Cover: 


19.8 


6.7 a (1.6) 


8.5 a (1.8) 


42.7 


19.7 a (4.3) 


18.5 a (3.9) 


Shrubs: 














Artemisia frigida 




0.3 


2 




1.4 


2.3 


Rosa arkansana 


2.7 


0.5 


0.2 


3.9 


1 


0.5 


Total Shrub: 


2.7 


0.8 a (1.2) 


2.2 a (1.3) 


3.9 


2.4 a (0.6) 


3.1 a (0.4) 


Introdlced Cover 


0.2 


0.6 a (0.5) 


0.9 a (0.4) 


3.6 


4.0 a (2.4) 


1.0 a (0.5) 


Richness (no71.5 m^) 


17.5 


23.3 a (1.8) 


24.3 a (1.3) 


21 


27.8 a (1.4) 


26.0 a (1.5) 


Shannon Diversity 


0.95 


1.08 a (0.04) 


1.08 a (0.04) 


0.99 


1.16 a (0.04) 


1.14 a (0.04) 


Other: 














Litter Cover 


11.2 


39.6 A (10.2) 


91.7 8(5.8) 


55.4 


76.3 A (4.1) 


97.9 B (1.7) 


Mulch Cover 


76 


52.1 A (8.0) 


7.5 B (5.6) 


28.2 


14.7 A (3.3) 


0.0 B (0) 


Bare Soil 


11.9 


5.4 a (3.6) 


0.1 a (0.1) 


15 


6.6 A (1.4) 


0.3 B (0.2) 


Exposed Rock 


1.3 


0.8 a (0.4) 


0.2 a (0.1) 


1.2 


0.5 a (0.3) 


0.1 a (0.03) 


Microphytic Cover 




2.9a(2.1) 


0.5 a (0.3) 




1.9 a (1.9) 


1.8a(1.7) 



*Within a component and year, cover values across the fire boundary with different upper and lowercase letters are signifi- 
cantly different from one another at p < 0.05 and p < 0. 10, respectively. 



56 



The Canadian Field-Naturalist 



Vol. 116 



Overall, the greatest amounts of bare soil and mulch 
were evident at the interior bum, further supporting 
the notion that this area was more severely burned. 

Potential for soil erosion was a serious concern to 
the local community and land administration agen- 
cies. Soil drifting was so severe on adjacent cropland 
immediately following the burn that emergency 
tillage was imposed. In contrast, there was little or 
no evidence of mineral soil loss on burned range- 
land. Soil exposure was less than 1% on unbumed 
areas, with 5.5 to 6.6% exposure on the perimeter 
burn and 11.9 to 15% for the interior burn during 
1998 and 1999, respectively. In Foothills Rough 
Fescue grasslands, soil erosion by water tends to 
increase when soil exposure exceeds 15% (Johnson 
1962; Naeth et al 1991); although soil exposure on 
the burned area increased, it failed to exceed this 
threshold. 

Comparative assessment of water quality above- 
and down-stream of the fire indicated that although 
total suspended and dissolved solids differed little, 
greater nitrogen was present in water collected from 
creeks immediately below the burned watersheds 
(Table 2; p = 0.051). Thus, fire likely contributed to 
nitrogen loss during snowmelt and/or spring rainfall 
from the watersheds. 

Herbage ANPP and Litter 

Graminoid and forb ANPP followed patterns simi- 
lar to that for plant cover (Table 3). In particular, 
graminoid production declined (p<0.05) by about 
40% across the fire boundary in 1998, but remained 
negligibly less in 1999, with full recovery in 2000. 
In contrast, forb production appeared to be greater 
on the burned area in the years after the fire (Table 
3), although this difference was not statistically sig- 
nificant (p>0.10). The greatest visible effect of 
burning in each year was at the interior bum, where 
a further decline in graminoid production and 
increase in forb production initially occured relative 
to the perimeter burn. Interestingly, this trend 
reversed in 2000, with graminoid and forb produc- 
tion apparently greater at the interior burn. 

The initial decrease in graminoid ANPP after fire 
is consistent with other studies for both Foothills 
Rough Fescue (Jourdonnais and Bedunah 1990) and 
Plains Rough Fescue grasslands (Redmann et al. 
1993; Gerling et al. 1995). Recovery of production 



in all these smdies took at least two years. Redmann 
(1978) attributed declines in production following 
buming to increased plant water stress. However, the 
favourable precipitation during 1998 in the current 
study suggests the reduction may be from plant 
responses directly caused by fire rather than a 
change in soil water regime. Coupland (1974, as 
cited in Redmann 1978) also documented reduced 
productivity after fire in Mixed Prairie despite 
above-normal precipitation. Regardless of the mech- 
anism, the magnitude of production decline observed 
here may be linked to the dormant season timing of 
the bum, as autumn fires are more detrimental than 
spring fires (Redmann et al. 1993). 

Several other factors may contribute to the reduc- 
tion in graminoid production. Although the plant 
communities examined were in good to excellent 
range condition at the time of the fire based on the 
composition of unburned areas, drought or heavy 
grazing prior to the fire may have increased stress on 
graminoids and reduced their vigor or winter hardi- 
ness. Additive negative effects on vegetation have 
been shown between defoliation and drought (e.g., 
Hendrickson and Berdahl 2002) and defoliation and 
fire (e.g.. Bunting et al. 1998). Although the com- 
bined impacts of defoliation and fire have been 
investigated on Foothills Rough Fescue (Bogen 
2001), that study examined defoliation after buming 
rather than before. 

The unusually mild autumn preceding the Granum 
fire may also have allowed vegetation to continue 
development into December. Erichsen-Arychuk et 
al. (2002) documented variable grassland recovery in 
Dry Mixed Prairie landscapes following August 
wildfire under drought conditions, with landscape- 
based differences potentially due to the stage of plant 
development at the time of fire. Following buming, 
the reduction of insulation through the loss of litter 
and associated snow cover would result in colder soil 
temperatures in winter (Johnston et al. 1971), 
increasing the susceptibility of burned plants to 
freezing (Kowalenko and Romo 1998). 

As expected, litter mass was markedly less 
(p < 0.05) on the bumed area (Table 3). Although the 
favourable growth during 1998 increased litter on 
the bum by the end of 1999, it remained less com- 
pared to the unburned area, with this trend continu- 



Table 2. Results of the analysis of water sampled from three creeks above and below the fire affected watersheds in 1998 
and analyzed for total suspended solids (TSS), total dissolved sediments (TDS), and nitrogen (N). 




Mean per Sampling 


Location (SE): 




Significance*: 


Variable: 


Above Burn (n = 11 ) 


Below Bum (n = 11) 


T- statistic 


Probability 


TSS (ppm) 
TDS (ppm) 

N (ppm) 


178.5(44.5) 
564 (56.4) 
1.15(0.15) 


297(152.4) 
1010(328.7) 
3.88(1.30) 




0.73 
1.42 

2.22 


p = 0.482 
p = 0.186 
p = 0.051 



*Paired t-tests contrast samples collected from above and below the bumed portions of the watershed. 



2002 



BoRK, Adams, Willms: Resilience of Foothills Rough Fescue 



57 



Table 3. Mean (SE) current annual graminoid and forb production, as well as litter levels (kg/ha) within the unbumed, 
perimeter bum, and interior bum locations in 1998, 1999, and 2000. 



Component 



Year 



Unbumed 



Perimeter Bum 



Interior Bum 



Graminoid 



Forb 



Litter 



1998 


1466(126) 


1999 


1596(154) 


2000 


966(109) 


1998 


439 (44) 


1999 


453 (37) 


2000 


110(15) 


1998 


898 (69) 


1999 


3351 (761) 


2000 


2101 (381) 



A* 



A 
A 



940 (77) B 


714 


1498 (226) 


1220 


844 (48) 


1349 


545 (73) 


682 


621 (105) 


921 


133 (29) 


292 


0(0) 





852 (98) B 


662 


908 (28) B 


948 



*Within a component and year, comparisons between bumed and unbumed means across the fire boundary represented by 
different letters are significandy different (p < 0.05). 



ing into 2000 (Table 3). Litter is important for con- 
serving soil water and maintaining herbage produc- 
tion (Weaver and Rowland 1952; Willms et al. 
1986). Antos et al. (1983) found the loss of litter 
after fire increased soil temperatures, reducing the 
near-surface effective water regime. Fluctuations in 
litter, even on the unbumed area (Table 3), indicate 
how rapidly this variable can change. Litter is lost as 
a result of fire, herbivory, microbial decomposition, 
and weathering, and is affected by grassland species 
composition (Facelli and Pickett 1991). Willms et al. 
(1996) found 23% of total biomass disappeared 
between fall and spring on good condition rough fes- 
cue grassland, while 56% disappeared from stands 
dominated by forbs and introduced grasses. Given 
the slow re-accumulation of litter in this study, it 
appears several years are needed for litter re-accu- 
mulation following the Granum fire. 

Rough Fescue Seedhead Production 

Burning increased (p<0.10) Rough Fescue seed- 
heads two years later, when seedhead densities on 
the burned area were more than twice that of the 
unbumed area (Table 4). The delay in response fol- 
lowing disturbance is similar to that found in other 
studies for Foothills Rough Fescue (Johnston and 
MacDonald 1967; Willms 1988). Gerling et al. 
(1995) found Plains Rough Fescue increased seed- 
head production the year following fire, but only 
when burned in spring or early summer (i.e., 1 June 
or earlier) of the previous year rather than in late 
summer or fall. 



Buming may have increased seedhead production 
by the addition of nutrients to the soil or the loss of 
litter. Litter removal may trigger seedhead establish- 
ment through increases in photosynthetically active 
radiation (PAR) within the meristematic region of 
the plant crown (Willms 1988). The delayed 
response indicates that either plant stress immediate- 
ly after buming inhibited reproduction the first year, 
or more likely, that there is a time lag needed for 
developing tillers to become reproductive, as deter- 
mined by the new environmental (e.g., light and/or 
nutrient) conditions after fire. It should also be noted 
that seedhead production was generally greater on 
the unbumed area in 1999 (19.5 ±3.8) than in 1998 
(2.9 ± 1.1) (Table 4). Thus, abundant seedhead pro- 
duction in 1999 on the burned area is due to the 
combined effect of favourable moisture during 1998, 
coupled with burning. The increase in seedheads 
reflects an important ecological adaptation of 
Foothills Rough Fescue to ensure this bunchgrass 
recolonizes areas of soil following buming, and thus, 
ensures its perpetuation within the plant community. 

Rough Fescue Forage Quality 

Burning positively affected the forage quality of 
Rough Fescue. In particular, crude protein and total 
digestibility increased, while acid detergent fiber 
(ADF) decreased (Table 5). However, between-plant 
variation was considerable, with only crude protein 
differing significantly (p<0.10) across the fire 
boundary in 1998 (other variables had p<0.15). 



Table 4. 


Rough Fescue seedhead production on the three sampling 


areas from 1998 to 2000. 








Seedhead Densities — #/m- 


(SE): 




Signi 


ficance*: 


Year 


Interior Burn 


Perimeter Bum 




Unbumed 


T- statistic 


Probability 


1998 
1999 
2()()0 


0.2(0.1) 

56.2(17.3) 

0.4(0.1) 


1.1(0.6) 

42.0(10.9) 

0.1 (0.1) 




2.9(1.1) 

19.5(3.8) 

0.8 (0.4) 


2.21 
2.64 
1.73 


p = 0. 1 1 
p = 0.08 
p = 0.18 



*Paired t-tests contrast data collected across the fire boundary (i.e. Perimeter Burn vs Unbumed). 



58 



The Canadian Field-Naturalist 



Vol. 116 



Table 5. Mean (SE) percent crude protein (CP), acid detergent fiber (ADF), and total digestible nutrients (TDN) of 
Foothills Rough Fescue plants sampled within the unbumed, perimeter bum, and interior bum locations in 1998 (n = 5 per 
transect) and 1999 (n = 8). 



Component 



Year 



Unbumed 



Perimeter Bum 



Interior Bum 



CP (%) 
ADF (%) 
TDN (%) 



1998 


7.20(0.11) 


1999 


7.39 (0.29) 


1998 


41.8(0.7) 


1999 


42.2 (0.4) 


1998 


52.1 (0.5) 


1999 


51.4(0.3) 



A 



A 



8.40 (0.44) b 


10.10 


6.54 (0.36) 


8.49 


40.2 (0.5) 


38.6 


40.9 (0.4) B 


38.9 


54.5 (0.3) 


56.9 


53.4 (0.3) B 


56.4 



*Within a nutrient, comparisons between bumed and unburned areas across the fire boundary represented by different 
uppercase (p< 0.05) or lowercase letters (p < 0.10) differ significantly. 



When larger sample sizes of plants were used in 
1999, energy and total digestibility were greater 
(p<0.05), and ADF lower (p<0.05), within the 
bumed area. Crude protein was unaffected (p > 0.10) 
in 1999, although it remained particularly high in 
plants sampled from the interior bum. 

The initial increases in protein are similar to the 
changes reported for Plains Rough Fescue following 
fire (Redmann et al. 1993) and burned Bluebunch 
Wheatgrass (Agropyron spicatum (Pursh) Scribn.) 
(Willms et al. 1981). Improved quality may be due 
to nutrient release into the soil following burning 
(Daubenmire 1968) or reduced competition on the 
burned area corresponding with a decline in live 
plant cover (Table 1). Increased quality may also 
stem from a delay in plant phenology throughout the 
summer, which would result in greater comparative 
quality at a fixed date of sampling. Delayed phenolo- 
gy of rough fescue grassland following autumn bum- 
ing was documented by Redmann et al. (1993), as 
was an associated increase in N concentration. 

Conclusions 

The 1997 Granum wildfire in Fescue Prairie had 
variable effects on Foothills Rough Fescue grassland 
plant species abundance and ground cover, herbage 
production, and forage quality, as determined by dif- 
ferences between the unbumed and perimeter burn 
locations. Grass cover and production declined the 
year after fire, but recovered by 1999, likely as a 
result of the above average precipitation in 1998. 
Drought after burning may have had an additive 
detrimental impact on vegetation recovery, particu- 
larly in the absence of litter to conserve soil water. 
Declines in production also coincided with increases 
in the forage quality of Rough Fescue. The observed 
responses highlight the impacts of dormant season 
wildfire on Foothills Rough Fescue grasslands, 
including the resilience of the dominant species. 
Additionally, it should be noted that although 
species richness and diversity were not increased by 
fire in this investigation, there was evidence that fire 
was effective in increasing the abundance of forbs, 
particularly legumes. Thus, the importance of occa- 



sional fire in maintaining biodiversity should not be 
discounted within these grasslands. 

For many ranches affected by the fire, the need to 
restore litter through conservative grazing may seem 
to conflict with their natural concem about grass litter 
as a fire hazard. Balancing the need for litter to con- 
serve water and protect the range resource, yet pre- 
vent excessive fuel loading, will be an ongoing ques- 
tion for land managers to address. Fortunately, the 
more enduring negative impacts of fire appear to be 
confined to a relatively small area. The results of this 
project should also be of interest to the managers of 
protected areas where grazing and the normal cycling 
of biomass by large herbivores and fire have been 
altered from their natural process on the landscape. 

Acknowledgments 

This research was funded by the Alberta Cattle 
Commission (ILO Grant# 98-0987), with additional 
support from the University of Alberta, AAFRD- 
Public Lands, AAFC-Lethbridge, and Norwest Labs. 
The assistance of Amanda Bogen, Darlene Moisey, 
Janna Wowk, and Dianne Nadeau with field setup 
and data collection is greatly appreciated. This 
research was made possible by the cooperation of 
many ranchers within the area affected by the 
Granum wildfire. Their interest and dedication is 
greatly appreciated. 

Documents Cited (marked * in text) 

Tymstra, C. 1998. The 1997 Granum fire in southwest 
Alberta: A case study. Unpublished Alberta Environ- 
mental Protection Report, Land and Forest Services, 
Forest Protection Branch. 10 pages. 

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Robberecht, R., and G. E. Defosse. 1995. The relative 
sensitivity of two bunchgrass species to fire. Inter- 
national Journal of Wildland Fire 5: 127-134. 

Strong, W. L. 1992. Ecoregion and ecodistricts of Alberta. 
Volume 1. Alberta Forestry, Lands, and Wildlife, Land 
Information Services, Edmonton. 77 pages. 

Tannas, C. 1990. Range survey of selected grazing leases 
in the Southern Porcupine Hills. Alberta Forestry, Lands 
and Wildlife. Public Lands Division Report. 125 pages. 

Weaver, J. E., and N. W. Rowland. 1952. Effects of nat- 
ural mulch on development, yield, and structure of 
native grassland. Botanical Gazette 114: 1-19. 

White, R. S., and P. O. Currie. 1983. Prescribed burning 
in the Northern Great Plains: Yield and cover responses 
of three forage species in the mixed grass prairie. 
Journal of Range Management 36: 179-183. 

Willms, W. D., A. W. Bailey, A. Mclean, and C. Kalnin. 
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Willms, W. D., S. Smoliak, and J. F. Dormaar. 1985. 
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Willms, W. D., S. Smoliak, and A. W. Bailey. 1986. 
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Received 23 October 2000 
Accepted 21 March 2002 



Survival, Fates, and Success of Transplanted Beavers, 
Castor canadensis, in Wyoming 

Mark C. McKinstry and Stanley H. Anderson 

Wyoming Cooperative Fish and Wildlife Research Unit', Box 3166, Laramie, Wyoming 82071 USA 

McKinstry, Mark C, and Stanley H. Anderson. 2002. Survival, fates, and success of transplanted Beaver, Castor canaden- 
sis, in Wyoming. Canadian Field-Naturahst 1 16(1): 60-68. 

Beaver {Castor canadensis) through their dam building activities, store water, trap sediment, subirrigate vegetation, and 
subsequently improve habitat for fish, wildlife, and livestock. Many landowners realize the benefits that Beaver can bring 
to a riparian area and are interested in using them to improve this habitat. From 1994 to 1999 we trapped and relocated 234 
Beaver to 14 areas throughout Wyoming to improve riparian habitat and create natural wetlands. We attached radio trans- 
mitters to 1 14 Beaver and subsequently determined movements and mortality of released Beaver, and the overall success of 
our releases. Mortality and emigration (including transmitter failure) accounted for the loss of 30% and 51%, respectively, 
of telemetered Beaver within 6 months of release. Kaplan-Meier survival estimates were 0.49 (SE = 0.068) for 180 days 
and 0.433 (SE = 0.084) for 360 days, and did not differ significantly between age classes. On average, 17 Beaver were 
transplanted to each release site, and at 1 1 locations, in an attempt to augment single Beaver that had become established 
and increase transplant success, we transplanted Beaver in two or more years. Success of an individual Beaver's relocation 
was unrelated to any of the variables we tested, although 2-3.5 year-old Beaver had higher average success (measured in 
days of occupancy at the release site) than older animals. Animals < 2 years old had 100% mortality and emigration losses 
within 6 months of release. High predation and mortality rates of our released Beaver may be due to habitat (our streams 
were shallow with no ponds and provided little protection) and extensive predator communities. We established Beaver at 
13/14 of our release sites and they eventually reproduced. Our results show that Beaver can be relocated successfully but 
losses from mortality and emigration need to be considered and planned for. 

Key Words: Beaver, Castor canadensis, transplanting, reintroduction, translocation, predation, Wyoming 



Beaver {Castor canadensis) alter riparian-stream 
ecosystems through their woodcutting and dam- 
building activities. Beaver dams create a lentic habi- 
tat in an otherwise lotic system. These ponds retain 
sediment and organic matter in the channel, create 
and maintain wetlands, modify nutrient cycling and 
decomposition dynamics, modify the structure and 
dynamics of the riparian zone, alter hydrologic 
regimes (Butler 1991), and influence the character of 
water and materials transported downstream 
(Naiman et al. 1988). The resultant habitats are rich 
mosaics of diversity that are beneficial hydrological- 
ly (Hanson and Cambell 1963; Rabe 1970; Johnson 
et al. 1992), biologically (Jenkins and Busher 1979 
reviewed by Hill 1982; Olson and Hubert 1994 
Brown et al. 1996; McKinstry and Anderson 1999 
McKinstry et al., 2001), and socially (Naiman et al. 
1988). It is estimated that Beaver have been extirpat- 
ed from over 25% of the streams in Wyoming, and 
in many streams where they are still present their 
numbers have been reduced to where they are eco- 
logically absent (McKinstry et al. 2001). The elimi- 



'The Unit is jointly supported by the University of 
Wyoming, Wyoming Game and Fish Department, U.S. 
Geological Survey-Biological Resources Division, and the 
Wildlife Management Institute. 



nation of Beaver from portions of its historic range 
has been cited as a major influence on the structure 
and patterns of vegetation in these systems (Neff 
1957; Barnes and Dibble 1986; Naiman et al. 1986; 
Nummi 1989; Kay 1994; Nolet et al. 1994). 

Throughout the intermountain west, interest has 
been expressed in improving riparian areas for 
wildlife, livestock, and humans (Apple et al. 1985; 
McKinstry and Anderson 1999; McKinstry et al. 
2001). Beaver, through their dam building activities, 
can increase water storage, reduce sedimentation, 
and improve vegetation communities (Naiman et al. 
1988), all of which are valuable to many landowners. 
Livestock are also attracted to Beaver-influenced 
areas for water, shade, and vegetation that remains 
green after upland forage has dried out. Forage pro- 
duction near these wetlands is often two to three 
times higher than comparable upland ranges (Apple 
et al. 1985; Chancy et al. 1991: 31). Many states 
have undertaken Beaver transplant programs to 
improve riparian areas (Smith 1980; Hill 1987; 
Butler 1991; Collins 1993; Vore 1993; McKinstry 
and Anderson 1997; McKinstry 2001) and managers 
with the Wyoming Game and Fish Department 
(WG&FD) decided to investigate the feasibility of a 
Beaver relocation program in areas where beaver 
have not recolonized due to isolation from dispersing 
populations and poor habitat conditions. 

In 1994 we initiated research to (1) document the 



60 



2002 



McKlNSTRY AND ANDERSON: TRANSPLANTED BeAVERS IN WYOMING 



61 



effects of Beaver on riparian areas in Wyoming 
(reported in McKinstry et al. 2000; 2001), (2) assess 
Beaver management concerns from both private and 
public landmanagers (reported in McKinstry and 
Anderson [1999]), and (3) evaluate a Beaver reintro- 
duction project for the purpose of wetland creation 
and riparian improvement. Our objectives in this 
paper are to examine survival, mortality, emigration, 
and success of Beaver translocated in Wyoming for 
the purpose of riparian restoration. 

Study Area 

Beaver were trapped at 33 various locations in 
Wyoming (described in McKinstry and Anderson 
1998) and translocated to 14 different P^-3'<^ order 
streams (< 0.283 m/sec) throughout Wyoming 
(Figure 1, Table 1). All drainages were walked a 
minimum of 3 km in both directions from the pro- 
posed release site to document any past or current 
Beaver activity. At 13 of the release sites, old Beaver 
sign (20-100 yrs) was present but we found no fresh 
activity. At the remaining site, Breteche Creek, prior 
Beaver activity was not found. All release sites had 
sufficient vegetation to support Beaver. Four streams 
were ephemeral and dry in early August each year 



that we checked them (1993-1999), the remainder 
were perennial and carried water year-round (Table 

1). 

Methods 

Beaver were trapped using snares and Hancock 
traps (McKinstry and Anderson 1998) from areas 
where they were causing damage to landowners (pri- 
marily irrigation conflicts) (30 sites) or where they 
were so plentiful that selective removal would not 
significantly impact the habitat (3 sites). All Beaver 
were trapped from colonies that were dam and lodge 
builders (creek Beaver) as opposed to bank-denning 
non-dam builders (river Beaver). We felt that these 
animals would be more likely to create the desired 
habitat. 

We began trapping after ice-off in the spring (usu- 
ally early to mid May) and concluded trapping by 
the second week in October, depending on snow and 
ice conditions. All traps were set between 1600 h 
and 1900 h each day and were checked by 1000 h 
the following morning to minimize the time that ani- 
mals spent in traps. On average we set 17 traps/ 
night/trapper; more than that and it was difficult to 
get them checked by 1000 h and find them in the 






'n 



o 

T3 



iF 




Wyominj; 




Figure 1. Beaver relocation sites in Wyoming. Letters correspond to release site locations described in Tables I and 2. 



62 



The Canadian Field-Naturalist 



Vol. 116 



Table L Latitude and longitude, dominant vegetation, and stream classifications (using Rosgen's [1994] classification) 
for Beaver release locations in Wyoming. Letters for release sites correspond with locations in Figure 1. 



Stream Name (nearest town) 



Latitude, Longitude 



Dominant Vegetation 



Stream Classification 



(A) Bear Gulch (Story) 

(B) Breteche Creek (Cody) 

(C) Bush Creeka (Shell) 

(D) Currant Creek (Rock Springs) 

(E) Deep Creek^ (Sundance) 

(F) Ennos Creek (Thermopolis) 

(G) Lake Creek^ (Saratoga) 



(H) Little Red Creek (Casper) 
(I) Prairie Dog Creek (Big Horn) 
(J) S. Pine Creek^ (Sundance) 



(K) Red Creek (Rock Springs) 



(L) Spring Creek (Centennial) 
(M) Trabing Creek (Big Horn) 



44°31'46"N, 106°53'11"W 
44°24'36"N, 109°23'28"W 
44°27'41"N, 107°37'30"W 
41°12'55"N, 109°22'20"W 
44°47'29"N, 104°20'41"W 
43°54'50"N, 108°54'14"W 
41°28'16"N, 106°38'20"W 



42°42'55"N, 106°23'48"W 
44°35'36"N, 106°54'30"W 
44°44'29"N, 104°20'30"W 



41°04'01"N, 109°02'30"W 



41°12'59"N, 106°07'44"W 
44°36'01"N, 106°58'59"W 



Aspen and cottonwood 


DA5 


Aspen and cottonwood 


A4 


Cottonwood 


A4 


Willow 


05 


Gambel's oak and aspen 


C5 


Willow 


B4 


Narrowleaf cottonwood 




(Populus angustifolia) 




and willow {Salix sp) 


B4 


Cottonwood 


F4 


Aspen and cottonwood 


A5 


Scrub oak and common 




chokecherry 




(Prunus virginiana) 


F6 


Engelmann spruce 




{Picea engelmannii) and 




cottonwood 


F5 


Willow 


F5 


Aspen and western hawthorn 




{Crataegus succulenta) 


A4 



^streams were ephemeral and dry by early August each year we checked (1993-1999) 



thick willows where we were trapping. Beaver were 
sexed through cloacal examinations (Larson and 
Taber 1980:163), weighed to determine age class 
(kit, yearling, sub adult, and adult), and ear tagged in 
both ears with small monel ear tags for identifica- 
tion. Beaver were held up to five days post-capture 
in a 2.2 by 3.1 m cage that allowed them free access 
to water; this was necessary since we only caught a 
few (< 3) animals each day and we wanted to move 
them as a group to the release site which was always 
> 160 km away. 

Forty-six Beaver (range 1 1-31 kg) were implanted 
with Advanced Telemetry Systems (ATS, Isanti, 
MN) model 17 internal telemetry transmitters using 
techniques described by Davis et al. (1984). 
Implanted Beaver were monitored for a minimum of 
24 hours post surgery prior to release (Davis et al. 
1984). Beginning in 1998, 67 Beaver (range 
10-25 kg) were outfitted either with tail collars (8 
Beaver) or transmitters mounted on the tail using 
modified ear-tag transmitters (59 Beaver) (Roth- 
meyer et al. 2002) in an attempt to reduce logistical 
problems associated with surgeries conducted in the 
field (e.g., cost, time, sterility concerns). All radio 
transmitters were equipped with 24-hour mortality 
sensors. Animals were released after 1500 hours in 
an attempt to decrease predation during daylight 
hours. 

Beaver were monitored for movements and mor- 
tality for 2 days after release and approximately 
every 2 to 4 weeks thereafter. Mortality dates were 
calculated as the mid-date between the date found 
and the date of last-live location. At all release sites. 



streams were walked (minimum of 5 km) and flown 
(minimum of 10 km) in both directions 3 months 
after release to determine if Beaver were established. 
In four instances non-transmittered Beaver became 
established, and walking the streams allowed us to 
look for evidence of tree cuttings or dam construc- 
tion and thus determine if non-telemetered Beaver 
were active. Since we were interested in using 
Beaver to improve habitat within 3 km of the release 
sites (usually headwater areas), we defined emigra- 
tion as Beaver that moved further than 3 km from the 
release site. Beaver moving > 5 km from the release 
sites were not monitored unless they built dams and 
lodges and remained stationary. 

Cause of mortality was determined through physi- 
cal examination of hair and scat samples found at the 
kill site (Moore et al. 1974), bite marks and subcuta- 
neous bleeding (indicates animal was alive when bit- 
ten) on Beaver carcasses, track marks (O'Gara 
1978), and, beginning in 1998, DNA analysis of hair 
and scat samples. Dr. Elizabeth Williams, 
Pathologist at the Wyoming State Veterinary Lab, 
performed lab work and necropsies. Tom Moore and 
Deedra Hawk, Forensic Supervisor and Research 
Associate, respectively, Wyoming Game and Fish 
Department, examined hair and scat samples. 

We used a z test (Jandel Scientific 1994) to test 
proportional differences in captures of males versus 
females. For estimates of survival we used Kaplan- 
Meier product limit estimators (as reviewed byj 
White and Garrott 1990) for both 360 and 180 day 
survival rates. We used the Kaplan-Meier approach] 
since our data was staggered entry and we had con- 



2002 



McKlNSTRY AND ANDERSON: TRANSPLANTED BeAVERS IN WYOMING 



63 





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siderable loss of individuals due to emigration or 
transmitter failure. To model success of transplants 
we used both logistic and multiple regression 
(White and Garrott 1990). For our logistic regres- 
sion models we coded each Beaver as either a for 
failure (emigration or mortality) or a 1 for success 
(lived > 6 months, constructed a dam and lodge, and 
had the opportunity to reproduce). For multiple 
regression models we used the length of success 
(remained within 3 km of release site) in days as our 
dependent variable. We used weight, sex, age class, 
season of release (spring: May 9-June 15) or fall 
August 15-October 10), year of release 
(1994-1999), and number of cohorts released con- 
currently with each Beaver as our covariates. For 
age class we plotted weights and found natural dis- 
tinctions in the three younger classes (kits, yearlings 
[1.0-1.5-year olds], and subadults [2.0-3.5-year 
olds]) and grouped all animals > 13.6 kg (4-year 
olds) as adults. Minitab (Minitab 2000), SigmaStat 
(Jandel Scientific 1994), and SAS (SAS Institute 
1991) were used for all analyses. 

Results 

Snares and Hancock traps were used to capture 
277 Beaver at 33 locations throughout Wyoming and 
we eventually transplanted 234 to the 14 release 
sites. The 43 remaining Beaver either died during 
trapping (n = 15), transporting (n = 13), or were lac- 
tating females (n = 15), which we released after cap- 
ture. Trapping mortality was 10% for Hancock traps 
and 5.3% for snares, and was not significantly differ- 
ent (z = -0.07, P = 0.94, df = 11). Mortality from 
traps was due to becoming entangled in snares 
(n = 11) and being killed by predators while 
restrained in snares and Hancock traps (n = 4). 
Trapping success during our five years of trapping 
was 11.1 trap nights/Beaver or 9.0% (the probability 
of an individual trap's capture). Average weight of 
animals captured was 16.2 kg (range 2.4-31 kg. SD 
= 6.11) and there was no difference in proportion of 
males (0.42) or females (0.58) captured (z = 1.23, P 
= 0.22, df = 122). We transmittered 63 (55.3%) 
females and 51 (44.7%) males. 

Of 1 14 Beaver trapped, equipped with a transmit- 
ter, and relocated, 34 (30%) died within 180 days 
(x= 43 days, SD = 37.4) of release (Table 2). 
Another 7 Beavers (36% total mortality) died prior 
to the failure of their transmitters (181-503 days). 
Coyotes {Canis latrans) were responsible for at least 
27% of all mortalities, followed by Black Bears 
(Ursus americanits) (10%), Grizzly Bears {Ursus 
horribilis) (10%), Mountain Lions i^Felis concolor) 
(2%), and humans (5%). Another 22% (of all mortal- 
ities) died of unidentified predators and the remain- 
ing 24% died from undetermined causes. Within 180 
days of release, 58 (51%) Beavers either emigrated 
> 10 km from the release sites and were not found 



64 



The Canadian Field-Naturalist 



Vol. 116 



again, or developed faulty transmitters, which made 
them impossible to relocate. 

The Kaplan-Meier survival estimates for all 
Beavers were 0.49 for 180 days and 0.433 for 360 
days and did not differ between age classes (Table 
3). Beavers that died lived for an average of 86 days 
(range 1-503 days, SD = 1 14.8) until death, however 
eight died within seven days of release. All Beavers 
(that we found), except one, died within 0.5 km of 
the release site (the exception was found 0.75 km 
upstream of the release site). 

Twenty-three (19%) Beavers lived > 180 days and 
eventually built dams and lodges in the drainages 
where they were released. Additionally, a minimum 
of 10 (actual number could not be determined since 
we did not retrap Beavers after release) of the other 
120 Beavers released without transmitters were also 
found with dams and lodges within 3 km of the 
release sites (some of these may have been Beaver 
with faulty transmitters, although we feel this was 
unlikely). We released an average of 17 Beaver at 
each of our sites in an attempt to get animals to 
establish. Beaver successfully established at 13 of 
the 14 release sites and, as of September 2001, the 
13 sites were still occupied. At the unsuccessful site 
Beavers had conflicts with irrigation structures (e.g., 
damming irrigation ditches) and they were removed. 

We were unable to identify any variables in our 
analyses that significantly influenced the probability 
of success for beaver relocations. P values were 
> 0.2 and R-squared values were < 0.10 for all vari- 
ables and models tested. The 2-3.5 year old age 
class had greater average occupancy at a site (Figure 
2), although this relationship was not significant 
(P= 0.225, df 3). All kits and yealings (n = 12) 
either died (n = 5) or emigrated (n = 7) from the 
release site prior to 108 days and none were 
observed constructing dams and lodges. 

Discussion 

Limited range of transmitters and transmitter fail- 
ure may have influenced the number of animals we 



found after release (Rothmeyer et al. 2002), subse- 
quently increasing the number of Beavers assumed 
to have emigrated. Advertised range of the internal 
transmitters was 0.5 km but we found that the range 
was usually limited to 200 m, and animals within 
dens were not located until we were within 50 m of 
the transmitter. Range of the tail-mount transmitters 
was better but never approached the 1-km advertised 
range. Walking up and downstream of the release 
site within 20 m of the creek bed was necessary to 
determine movements and mortality. 

Animals emigrating or not found after release may 
have had higher survival rates but we believe this is 
improbable. More likely, these Beaver were killed 
and cached in holes and dens, or dragged out of 
range of receivers. Animals moving out of the vicini- 
ty of the release area may also have experienced 
higher predation rates due to increased exposure 
time and less time spent hiding in dense vegetation 
or constructing dens. Beavers, sympatric with Black 
Bears on islands in Lake Superior traveled shorter 
distances from water than those found on islands 
where bears were not present, posssibly a direct 
attempt to avoid predation (Smith et al. 1994). 
Survival estimates for unexploited (i.e. untrapped) 
adult beaver are generally 0.80 (Boyce 1974; 
Bergerud and Miller 1977; Bishir et al. 1983) but 
these animals have ponds and lodges for escape. 
High mortality (5/10) and loss (4/10) (never relocat- 
ed) rates were reported for beaver translocated in the 
James Bay area of Quebec, Canada (Courcelles and 
Nault 1983). Translocated animals undoubtedly have 
a higher susceptibility to predation for many reasons 
including unfamiliarity with the surrounding habitat, 
reduced fitness due to trapping stress, and possible 
exposure to more numerous and greater varieties of 
predators (Griffith et al. 1989; Stanley-Price 1989). 
Beaver have many natural predators and do not 
avoid predation through fighting, preferring instead 
to use water as an escape medium (as reviewed in 
Smith et al. 1994). Without ponds and dens to use 
for escape, Beaver are vulnerable to predation. 



Table 3. Kaplan-Meier 180 and 360 day survival estimates, sample sizes, 95% CIs, and SEs for Beavers translocated in 
Wyoming. Survival estimates were not computed for Beaver < 2 years old since 100% either died or emigrated prior to 180 
days. 



Age 
Class 



N 
Total 



n 
Surviving 



n 
Dyin^ 



n 
Unknown 



Survival 
Estimate 



95% CI 



SE 



180 days 

All ages 



2.5-3.5 years old 
4+ years old 



360 days 

All ages 
2.5-3.5 years old 
4+ years old 



114 
52 
51 



23 
13 
10 



34 
20 



58 
19 

33 



0.493 
0.436 
0.658 



0.358-0.628 
0.271-0.602 
0.449-0.868 



0.068 
0.085 
0.107 



114 
52 
51 



13 
7 
6 



36 

22 
8 



66 

23 
37 



0.433 
0.353 
0.658 



0.268-0.598 
0.167-0.538 
0.449-0.868 



0.084 
0.095 
0.107 



"animals either emigrated > 5 km or transmitters failed 



2002 



McKlNSTRY AND ANDERSON: TRANSPLANTED BEAVERS IN WYOMING 



65 



600 



500 

J. 

(0 400 

S 300 
0) 

o 
o 

3 200 
CO 

100 H 



1 



28 (6) 



"WT 



X 



89 (52) 



X. 



54(51) 



Kits Yearlings Subadults (2.0- Adults 

( 1 . 5 yrs old) 3 . 5 yrs old) (> 3 . 5 yrs old) 

Age Class 

Figure 2. Average survival, in days (n), for four age classes of relocated beavers. Error bars 
depict 95% CI's. 



Naturally dispersing Beaver have been seen in 
upland areas > 1 km from water sources and have 
been observed crossing mountain passes (Smith 
1980) far from water where they would be extremely 
vulnerable to predation. Movements across upland 
areas are undoubtedly successful but we question the 
frequency of this success in areas like Wyoming 
where multiple predators occur sympatrically. 

Translocated Beaver in North Dakota, Wisconsin, 
and Maine moved an average of 14.6, 7.4 and 11.2 
stream km, respectively, although the longest move- 
ment was 238 km (Hibbard 1958; Knudsen and Hale 
1965; Hill 1982) and naturally dispersing Beaver in 
Idaho and Quebec moved an average 8.5 and 18 km 
(Leege 1968; Courcelles and Nault 1983). While 
movements greater than 3 km are common for 
Beaver, exposure rates certainly increase during long 
movements through non-ponded habitat. Our ani- 
mals may have been attempting to return to trapping 
locations, which were greater than 160 km away. 
Since our goal was to establish Beavers in uncolo- 
nized areas, predation risk was unavoidable and high 
predation and emigration losses should be consid- 
ered when planning releases. 

Wildlife managers in Europe have made several 
attempts to transplant Beaver (C. fiber) with varied 
success (Zurowski and Kasperczyk 1988; Hartman 
1994; Nolet and Baveco 1996). Initially, mortality 
and emigration rates were high (14-36% and 23%, 
respectively) and were greatest in juvenile and year- 
ling animals. Mortality rates declined with the estab- 



lishment of dens and dams and populations became 
well established over time. However, transplanting 
in Europe usually occurs in major river systems 
where flooding of the dens is the primary disruptive 
event and predator populations are not as large 
(Nolet and Baveco 1996). In Wyoming, predator 
communities are well developed and are thought to 
be increasing (McKinstry and Anderson in review). 
Additionally, Beaver introduced in large rivers 
(> 2.83 m/sec) have greater aquatic escape cover 
than Beaver released in the small (< 0.283 m/sec) 
streams where we were working. 

The value of riparian areas for wildlife has been 
emphasized by many authors (as reviewed by 
Naiman et al. 1988), but predator-prey relationships 
in riparian areas and how they relate to habitat quality 
and availability are not well understood (Smith et al. 
1994). During drought periods animals may concen- 
trate in riparian areas along with their respective 
predators. These predators, while normally dependent 
upon another prey species, may find Beaver easy 
prey. Many wildlife species use riparian areas more 
frequently during fall due to increased forage and 
water in the riparian areas. Predators may also con- 
centrate in these areas to take advantage of higher 
prey densities, greater water availability, and lowered 
temperatures. Our releases were primarily (71%) 
done during the fall (15 August- 10 October) to take 
advantage of the Beaver's natural tendency to begin 
construction of dams and lodges in the fall (Vorc 
1993). We did spring releases to establish Beaver in 



66 



The Canadian Field-Naturalist 



Vol. 116 



ephemeral streams that were normally dry by mid 
August and to supplement one or two Beaver that had 
become established the year before. Relocations in 
the spring may have lower predation risks, although 
we did not see this relationship in our analyses. 

Several biologists suggested that we create small 
ponds at release sites to provide temporary refuge for 
released Beaver. In the two instances (Bear Gulch 
and Spring Creek; see Table 2 for specific numbers 
released) where we released Beaver into remnant 
ponds they emigrated from those areas within 10 
days and constructed their own dams and lodges 
elsewhere in the drainage, although the remnant 
ponds may have provided them with initial protec- 
tion from predators and helped to acclimate them to 
the area. Furthermore, creating small ponds is cost 
prohibitive and unlikely to be used in future trans- 
plants, therefore we did not consider this as a legiti- 
mate tool in our transplants. 

Our goal was to establish Beaver at release sites, 
therefore we continued to release Beaver until dams 
and lodges were constructed. On average we 
released 17 Beaver/site before they constructed a 
dam and lodge and successfully reproduced. In 11 
instances a single Beaver created a pond with a 
lodge and we transplanted additional animals in the 
hope that they would pair-up. We expected that 
many Beaver would emigrate from our release sites 
in efforts to return home, search for mates, or look 
for more suitable habitat. High predation and emi- 
gration rates for introduced Beaver should be expect- 
ed (Griffith et al. 1989) and planned for in any 
Beaver translocation project. 

We found no significant predictors of success or 
survival in our analyses. The 2-2.5 year-old Beavers 
had greater average success (Figure 2) and other 
researchers (Vore 1993; P. Jensen, Pennsylvania 
State University, personal communication) have sug- 
gested that this age class may be more suitable for 
recolonizing new areas since they are predisposed to 
emigrating, establishing new territories, constructing 
dams and lodges, and attracting mates. Our results 
are inconclusive but we recommend translocating 
Beavers > 2 years old since mortality and emigration 
of our younger animals totaled 100% within six 
months. 

Our results also call attention to programs where 
only one or two Beaver are relocated to either unoc- 
cupied habitat, usually under the pretext of improv- 
ing habitat, or to areas with existing Beaver popula- 
tions. These activities are common in the western US 
and are usually carried out by district biologists or 
conservation officers to remove nuisance Beaver 
without using lethal techniques. We suggest that 
unless managers are committed to successfully intro- 
ducing Beaver through planned introductions, moni- 
toring, and follow-up releases their time and money 
may be better spent on other duties. 



In summary, we found that Beaver could be used 
to create natural wetlands and improve riparian habi- 
tat in 13/14 streams where we relocated them. 
Mortality (30%) and emigration (51%) totaled 81% 
within the first 6 months of release and we needed to 
relocate an average of 17 Beaver/site to get a pair to 
establish and reproduce. Additional releases were 
also needed to augment single animals that had 
become estabhshed. Wildlife response to our created 
habitats was immediate (McKinstry et al. 2001) and 
landmanagers found that the habitats were valuable 
to both wildlife and livestock. We caution that 
releases should only be used in drainages where con- 
flicts with irrigation or road crossing structures are 
minimal and preferably where the drainage is con- 
trolled by a few landowners to simplify evaluation 
and management. 

Acknowledgments 

This research was supported by grants from the 
Wyoming Game and Fish Department, U.S. Fish and 
Wildlife Service, Rocky Mountain Elk Foundation, 
Ducks Unlimited, National Rifle Association, 
National Fish and Wildlife Foundation, North 
American Wetlands Conservation Council, Jack's 
Plastic Welding, George E. Menkins Memorial 
Fellowship, and the University of Wyoming. Field 
assistance, provided by Rory Karhu, E. Hardgrave, 
S. Rothmeyer, S. Mohren, and K. Rompola was 
much appreciated. Our thanks are also extended to 
the Breteche Creek Foundation, T. Malmberg, M. 
Miller, U.S. Forest Service, US Bureau of Land 
Management, the State of Wyoming, and numerous 
other landowners for allowing us access to release 
sites. L. Apple, J. Vore, K. Gordon, G. Butler, and 
two anonymous reviewers suggested revisions that 
improved the manuscript. 

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Received 1 November 2000 
Accepted 18 March 2002 



Songbird Community Composition Versus Forest Rotation Age in 
Saskatchewan Boreal Mixedwood Forest 

Enid E. Gumming ^ and Antony W. Diamond^ 



'1542 Empress Avenue, Saskatoon, Saskatchewan S7K 3G3 Canada 

^Atlantic Co-operative Wildlife Ecology Research Network, University of New Brunswick, P.O. Box 45111, Fredericton, 
New Brunswick E3B 6E1 Canada 

Camming, Enid E., and Antony W. Diamond. 2002. Songbird community composition verses forest rotation age in 
Saskatchewan boreal mixedwood forest. Canadian Field-Naturalist 116(1): 69-75. 

Songbird communities were characterised using unlimited-distance point counts in four age-classes of boreal mixedwood 
forest in central Saskatchewan in 1991. Forest age-classes surveyed were: mature stands (50-60 years old), rotation age 
stands (80-90 years), old stands (100-1 10 years), and very old stands (>140 years). Ten species (Winter Wren, Troglodytes 
troglodytes; Golden-crowned Kinglet, Regulus satrapa; Ruby-crowned Kinglet, R. calendula; Swainson's Thrush, 
Catharus ustulatus; Tennessee Warbler, Vermivora peregrina; Blackbumian Warbler, Dendroica fiisca; Magnolia Warbler, 
D. magnolia; Bay-breasted Warbler, D. castanea; Rose-breasted Grosbeak, Pheucticus ludovicianus and Evening 
Grosbeak, Coccothraustes vespertinus, were significantly more abundant in forest that was older than rotation age. Four of 
these species (Winter Wren, Tennessee, Magnolia and Bay-breasted warblers), reached their highest densities only in the 
oldest (>140) age class. Two species (Red-eyed Vireo, Vireo olivaceus, and Ovenbird, Seiurus aurocapillus) were signifi- 
cantly more abundant in forest that was rotation age or younger. In boreal mixedwood forest, habitat-selection patterns of 
songbirds suggest that many species, not also occurring in rotation-aged or younger stands, may require substantially older 
forest in order to persist. 

Key Words: songbirds, boreal, mixedwood forest, rotation age. 

Boreal forest is one of the largest terrestrial ecosys- 
tems on the planet (Shugart et al. 1992; Apps et al. 
1995). It comprises 20% of the world's forest and is 
the largest terrestrial ecozone in Canada, covering 
more than one-third of the country (Rowe 1972; 
Shugart et al. 1992). The southern part of this forest, 
the boreal mixedwood, supports the best tree growth 
(Rowe 1972) and is under the most pressure from 
human activities. In the three Prairie Provinces, more 
than 90% of the boreal mixedwood forest has been 
leased for harvesting under Forest Management 
Agreements (Cummings et al. 1994; Pratt and Urqhart 
1994; Alberta Environmental Protection 1998*). In 
Saskatchewan, over 80% of trees harvested come 
from mixedwood forest, predominantly White Spruce 
(Picea glauca) and Trembling Aspen {Populus tremu- 
loides), especially from the older age classes (Spencer 
1993). This type and age of forest accounts for, at 
most, 10% of the commercial forest (Kabzems et al. 
1 986), yet it is also the most biologically diverse habi- 
tat in the region (Welsh 1981; Harris et al. 1984; 
Acton et al. 1998). Thus, the biologically richest 
stands are being harvested in a proportion far exceed- 
ing their abundance (Erskine 1977). 

Canada is required under the Biodiversity Con- 
vention to ensure the ecological sustainability of all 
its ecosystems (Biodiversity Science Assessment 
Team 1994*), and the global market is demanding 
that suppliers of forest products demonstrate the 
environmental sustainability of their harvesting prac- 
tices (Canada 1995*). As boreal mixedwood forest 



comes under increasing pressure from harvesting, 
foresters will be expected to maintain biodiversity 
(CCFM 1992*; Freedman et al. 1994; Middleton 
1994*) but currently lack the information required to 
do so. When a previously little-exploited segment of 
the forest resource, such as old-growth mixedwood 
boreal stands, becomes the focus of intensified har- 
vesting, the assessment of impacts on biodiversity 
becomes an urgent priority. 

Our study was designed to address the question: 
"Are there elements of the biological community of 
old-growth boreal mixedwood forest whose survival 
may be threatened by harvesting?". In his 1977 study 
on boreal songbirds, Erskine suggested that the 
"...mature stage of the most fertile needle-leafed 
community" is biologically the most diverse commu- 
nity in the boreal forest and "...deserves more atten- 
tion from environmental impact studies than they 
have received". Specifically, we used songbirds as 
focal organisms, expecting that they might be useful 
indicators of impacts on other organisms, and per- 
haps of biodiversity as a whole (Freedman et al. 
1994). Songbirds are primarily insectivorous, feed- 
ing sufficiently close to the top of their food-chains 
to be potentially useful indicators of impacts on a 
wide range of organisms (Diamond and Filion 1987; 
Furness and Greenwood 1993). 

We chose a range of stand ages around the pro- 
posed rotation age (80-90 years), to assess whether, 
in the extreme case of all stands older than rotation 
age being eliminated from the landscape, an impact 



69 



70 



The Canadian Field-Naturalist 



Vol. 116 



Table \. Species which occurred significantly more or less frequently in a given forest age (Anova with 
Tukey's test). 



Species 


F 


DF 


P 


Significant Age 


Winter Wren 


87.1 


3 


*** 


140 > 


Tennessee Warbler 


3.0 


3 


* 


140 > 


Magnolia Warbler 


4.8 


3 


** 


140 > 


Bay-breasted Warbler 


6.9 


3 


*** 


140 > 


Ruby-crowned Kinglet 


6.1 


3 


** 


100 > 


Blackbumian Warbler 


7.1 


3 


*** 


100 > 


Black-throated Green Warbler 


4.3 


3 


** 


100 > 


Rose-breasted Grosbeak 


4.5 


3 


** 


100 > 


Swainson's Thrush 


7.2 


3 


*** 


50 < 


Red-eyed Vireo 


4.3 


3 


** 


80 > 


Ovenbird 


12.6 


3 


*** 


140 < 



*<0.05, **<0.01, ***<0.001 



on the bird community could be expected. Our 
youngest age-class was 30 years younger than rota- 
tion age. We did not study stands younger than this, 
because other studies have shown the songbird com- 
munity to be very different between young (<30) and 
old (>80) forest (Titterington et al. 1979; Schick et 
al. 1995; Hobson and Bayne 1999). In addition, 
intensified management will lead to an increased 
abundance of younger age-classes at the expense of 
those older than rotation age (Spencer 1993; 
Weyerhauser Canada 1998*). 

It is already known that a reduction in stand age 
may have negative effects on some bird species 
(Titterington et al. 1979; Welsh 1981; Farr 1993; 
Schick et al. 1995), but effects on particular species 
in spruce-dominated mixedwood forest have not pre- 
viously been addressed. Previous studies have shown 
that old mixedwood forests have a rich assemblage of 
bird species (Erskine 1977; Kirk et al. 1996). Many 
of these species do not occur in younger forest, or in 
hardwood dominated forest, or do so only at low den- 
sities (Stelfox 1995; Hobson and Bayne 1999). 
Methods 

This study was carried out in summer 1991 in and 
around Prince Albert National Park, Saskatchewan 
(53° 35'N 106° OO'W) (see Bayne and Hobson 1997), 
in boreal mixedwood forest (Kabzems et al. 1986; 
Acton et al. 1998). Ten study sites were established in 
forest of four different ages: three stands >140 years 
old, three 100-1 10 years old, two 80-90 years old and 
two 50-60 years old, and were aged by coring six of 
the largest trees on each site. All sites were mixed- 
wood forest and were as similar as field conditions 
allowed. Sites were pre-selected for similarity using 
Saskatchewan forest inventory maps, which are based 
on similarity of tree density and species composition 
(Kabzems et al 1986). Ground truthing showed stands 
were similar in species composition and tree density, 
however, older sites did tend toward greater abun- 
dance of conifers (Cumming 1995). The largest and 
most abundant trees in all stands, were White Spruce 



and Trembling Aspen. Two other species. White 
Birch (Betula papyrifera) and Balsam Fir {Abies bal- 
samea) also occurred (Cumming 1995). The main 
shrubby species were young Balsam Fir, White Birch, 
Alder {Alnus sp.) and Beaked Hazel {Corylus comu- 
ta). Ground cover was mainly composed of several 
species of moss; Bunchberry {Cornus canadensis); 
Twinflower (Linnaea borealis) and Wild Sarsaparilla 
(Aralia nudicaulis). 

Birds were counted using the unlimited-distance 
point count method (Bibby et al. 1992), which is 
widely used as the method of choice when large 
areas need to be surveyed rapidly and efficiently 
(Vemer 1985; Farr 1993; Bancroft et al. 1995). The 
method is a good indicator of relative abundance 
(Vemer and Ritter 1985; Bancroft et al. 1995) and an 
efficient method for comparing species composition 
and richness between habitats (Blondel et al. 1981). 
Point counts were at least 200 m apart and 100 m 
from a stand edge; these distances minimise the 
chances of double-counting, and including species 
from other habitats, respectively. Surveys were con- 
ducted between 04:00 and 09:00, by the same 
observer (EEC), and included all birds seen or heard 
in 10 minutes. Each site was surveyed twice during 
the breeding season; early (1-14 June) and late 
(20 June-4 July). Such spacing of surveys gives a 
more representative picture of a bird community 
than either one survey, or two carried out on consec- 
utive days (Hilden 1981; Skirven 1981). 

Because sampling effort differed between stands, 
(five point count stations/site in the oldest category, 
and three/site in the other three categories), the data 
were standardised by dividing the maximum number 
of individuals of each species recorded in a particu- 
lar age class by the number of point counts. This 
gives the probability of detection for a particular 
species in that age of forest. Birds that were flying 
overhead, or had very large territories, such as rap- 
tors, ravens, and woodpeckers were not included in 
the analysis. 



2002 



Gumming and Diamond: Songbird Community Composition 



71 



Bird Species 



Winter Wren 
Cape May Warbler 
Western Tanager 
White- winged Crossbill 
Gray Jay 

Golden-crowned Kinglet 
Rose-breasted Grosbeak 
Evening Grosbeak 
Brown Creeper 
Boreal Chickadee 
Ruby-crowned Kinglet 
Red-breasted Nuthatch 
Solitary Vireo 
Yellow-rumped Warbler 
Mourning Warbler 
Chipping Sparrow 
White-throated Sparrow 
Pine Siskin 
Swainson's Thrush 
Tennessee Warbler 
Blackbumian Warbler 
Black-throated Green Warbler 
Magnoha Warbler 
Bay-breasted Warbler 
Red-eyed Vireo 
Ovenbird 



50-60 



Forest Age (years) 
80-90 100-110 



<0.20 

0.20 to 0.49 
0.50 to 0.79 
0.80 to 1.00 
>1.00 



140+ 





Number of Species 



15 



22 



26 



* 



** 



more abundant in post rotation age sites (p<0.05) 



* 






* 




* 
* 

** 
** 



more abundant in sites at or below rotation age (p<0.05) 
Figure 1 . Probability of detecting a given species of bird at a point count in the different forest age classes. 



! 



Vegetation was measured in 0.04 ha circles (22 
metres in diameter), using a modified James and 
Shugart (1970) method. Two such circles were locat- 
ed near each point count station, one centred on the 
point count location, with the other located in a ran- 
dom direction 50 metres away. In each plot, the 
species, diameter (dbh), and height of each tree was 
recorded. Plants with a dbh greater than 7 cm were 
considered trees, while those less than 7 cm dbh 
were classified as shrubs (James and Shugart 1970). 
Percent shrub cover was estimated for the entire plot, 
and ground cover was estimated in four randomly 
located 1 m- quadrats. 



Data were analysed using 1-way Anova (Zar 
1996) using the statistical package SPSS. In the case 
where more than two categories were used, a 
Tukey's HSD test was applied to find where differ- 
ences had occurred (Zar 1996). Both bird and vege- 
tation data were analysed using four age classes and 
by pooling data for two age classes, either post-rota- 
tion age or rotation age and younger. 

Results 

The number of songbird species recorded in each 
habitat progressively increased with age, from 
youngest to oldest the numbers of species were: 15. 



72 



The Canadian Field-Naturalist 



Vol. 116 



Table 2. Species which are found significantly more often 
in post-rotation age forest. 



Species 


F 


DF 


P 


Winter Wren 


15.8 




*** 


Golden-crowned Kinglet 


3.9 




* 


Ruby-crowned Kinglet 


5.0 




* 


Swainson's Thrush 


5.3 




* 


Tennessee Warbler 


4.1 




* 


Magnolia Warbler 


6.1 




* 


Blackbumian Warbler 


8.7 




*** 


Bay-breasted Warbler 


5.9 




* 


Rose-breasted Grosbeak 


4.7 




* 


Evening Grosbeak 


9.6 




** 


Red-eyed Vireo^ 


6.8 




** 


Ovenbird" 


7.1 




** 



*<0.05, **<0.01, ***<0.001 

tOvenbird and Red-eyed Vireo were found significantly 
more often in forest of rotation age and younger. 



18, 22 and 26 (Figure 1). Eleven species were not 
detected in the youngest (50-60 years) forest sam- 
pled, but no species detected in the youngest forest 
did not also occur in the oldest (Figure 1). 

A 1-way Anova with Tukey's test found that 
Winter Wren, Tennessee, Magnolia, and Bay- 
breasted warblers were significantly more abundant 
in the >140 year old forest (Table 1). Ruby-crowned 
Kinglet, Rose-breasted Grosbeak, Blackburnian and 
Black-throated Green warblers (Dendroica virens) 
were most abundant in the 100-110 year old forest 
while Red-eyed Vireos were most abundant in the 
80-90 year old forest. Swainson's Thrushes were 
least abundant in the 50-60 year old forest, and 
Ovenbirds were least abundant in the >140 year old 
forest (Table 1). Pooling the data into pre and post- 
rotation age showed ten species to be significantly 
more abundant in post-rotation aged forest (Table 
2). Only Ovenbird and Red-eyed Vireo were more 
abundant in forest of rotation age and younger 
(Table 2). 

The percent of coniferous trees relative to decidu- 
ous trees increased with increasing forest age (Table 
3). The average height of the forest was not signifi- 
cantly different between the three oldest ages, but 



the average tree height of the 50-60 year old forest 
was significantly shorter than the other three (Table 
3). The amount of shrub cover and moss cover was 
significantly higher in the oldest forest, while the 
amount of litter was significantly higher in the 
youngest forest (Table 3). The average height of the 
combined post-rotation age forest (20.4 ± 7.0 m) 
was significantly taller than the rotation age and 
younger forest (15.8 ± 3.9 m) (F = 30.0, DF = 1, 
P< 0.001). 

Discussion 

All bird species found in the youngest stands also 
occurred in older forest; however, the reverse was 
not true. Species were added to the forest with 
increasing age, and species richness increased with 
stand age. We sampled more old than young stands, 
therefore, with increased sample effort some of the 
missing species may indeed be detected in younger 
forest. However, our data indicate a definite trend 
for higher species richness in forest that is older 
than rotation age and for many species there is also 
a trend for greater abundance in forest older than 
rotation age. This may be caused by old forest hav- 
ing higher habitat heterogeneity than young forest, 
which is thought to contribute to its increased 
species richness ( Meslow et al. 1981; Hunter 
1990). This increased heterogeneity has a variety of 
causes, including windthrow gaps, and tree disease 
and death, which allows for infilling with various 
ages and species of trees (Erskine 1977; Hunter 
1990). 

Studies in Maine, Ontario, and Alberta found 
Ruby and Golden-crowned kinglets. Winter Wrens, 
crossbills (Loxia spp.). Cape May, Blackbumian, and 
Bay-breasted warblers were more abundant in 
mature to old forest and present at either much lower 
densities, or absent, in younger stands (Titterington 
et al. 1979; Welsh 1981; Schieck et al. 1995). A 
small sample size was the most likely reason we did 
not find Cape May Warbler, Western Tanager 
{Piranga ludoviciana) and White-winged Crossbill 
{Loxia leucoptera) significantly correlated with post- 
rotation aged forest. However, as they were observed 
only in the oldest forest sampled, this suggests a 



Table 3. Percent of coniferous and deciduous tree cover and average height at different ages, and percent of shrub and 
ground cover at different ages. Percentages are mean ± 1 standard deviation. 





% 


% 






Tall Shrub 








Ground Cover 






























Age 


Conifer 


Decid. 


Height (m) 


% Cover 


% 


Moss 


% 


Herb 


% Litter 


% Grass 


140+ 


80 


20 


20.1 


±7.5 


41.7 


± 24.6* 


53.0 


±28.3* 


4.3 


±3.5 


42.8 ± 


27.2 







100-110 


65 


35 


21.1 


±6.1 


28.8 


±22.3 


9.4 


±17.4 


22.5 


±16.5 


65.6 ± 


20.9 


2.5 ± 


7.1 


80-90 


48 


52 


18.3 


±4.3 


22.5 


±38.6 


2.5 


±5.0 


25.0 


±28.9 


70.0 ± 


29.4 


2.5 ± 


5.0 


50-60 


49 


51 


13.9 


±2.0* 


2.5 


±3.9 


3.1 


±2.4 


4.4 


±2.3 


92.5 ± 


12.9* 








*Indicates means that are significantly different at P<0.05; 1-way Anova;Tukey's HSD. 



2002 



Gumming and Diamond: Songbird Community Composition 



73 



trend for these species to use mature to old forest, 
especially as they have been found to be significant- 
ly associated with older forest in other studies (Farr 
1993; Benkman, 1993). 

Some species did not show a significant difference 
in their habitat preference and occurred at similar 
densities in all ages tested. Species such as, Solitary 
Vireo {Vireo solitarius); Yellow-rumped Warbler 
(Dendroica coronata); Chipping Sparrow (Spizella 
passerina) and Pine Siskin (Carduelis pinus), have 
been found to be associated with coniferous forest, 
including Jack Pine (Pinus banksiana), rather than 
forest age per se. (Kirk et al. 1996); and species such 
as White-throated Sparrow {Zonotrichia albicolliss), 
are widely distributed across a great many forest 
types and ages (Kirk et al. 1996; Hobson and Bayne 
1999). 

Many of the 26 species sampled showed some 
degree of dependence on forest as old as, or older 
than, the rotation age of 80-90 years. Eight species 
(30% of the total community) occurred exclusively 
in stands substantially older than rotation age, and 
three more occurred only in these stands or those of 
rotation age; thus 42% of the species in this commu- 
nity showed a marked degree of dependence on 
mature or old stands. In addition, five other species 
(Swainson's Thrush, Tennessee, Blackburnian, 
Magnolia, and Bay-breasted warblers), that occurred 
in all age classes, were significantly more abundant 
in the post-rotation age forest. We conclude that 
62% of these species (16/26) are likely to suffer 
some reduction in population density if stands older 
than rotation age are altogether lost from the land- 
scape. 

Several of the species which occurred in younger 
stands did so at low abundances; for these species, 
young stands could potentially represent population 
sinks (van Home 1983; Pulliam 1988), in which 
populations can only be maintained by immigration. 
Many of the species that were age-sensitive also 
appear to be area-sensitive (Hobson and Bayne 
2000). As these species appear to show sensitivity to 
age and fragmentation, we suggest that there is a 
need for further research on the habitat requirements 
and reproductive biology of these species. This is 
especially true as many of these species may also be 
facing habitat loss in the southern fringe of the bore- 
al mixedwood forest (Cumming et al. 2001; Hobson 
et al. in press). 

We conclude that managed boreal mixedwood 
forest should maintain some stands of at least 120 
years or older if the communities characteristic of 
the whole forest ecosystem are to be sustained. 
Natural disturbance patterns could be used to guide 
the proportion of the landscape to be retained in 
this age-class. The question of the most appropriate 
size and configuration of such old-growth stands, in 
relation to conservation of songbirds, was not 



addressed by our study, but we draw attention to 
abundant evidence that reduction in forest age, and 
fragmentation of forest patches (a process that has 
accelerated in central Saskatchewan in recent years, 
Fitzsimmons et al. 1997*) can have serious impli- 
cations for the sustainability of songbird popula- 
tions (Schieck et al. 1995; Villard et al 1995; 
Cumming et al. 2001) 

Acknowledgments 

This study was carried out in partial fulfilment of 
the requirements for the M.Sc degree at the 
University of Saskatchewan by E. E. Cumming. 
Parks Canada and Weyerhauser Canada Ltd. allowed 
field work to be carried out in Prince National Park 
and on Weyerhauser lease, respectively. Funding for 
this project was provided by the Canadian Wildlife 
Service (which also provided equipment and office 
facilities at the Prairie and Northern Wildlife 
Research Centre), Prince Albert National Park, 
Nature Saskatchewan and the Saskatchewan Wildlife 
Habitat Development Fund. This manuscript was 
improved by helpful comments by A. J. Erskine, K. 
A. Hobson, S. L. Van Wilgenburg and an anony- 
mous reviewer. 

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Biodiversity Science Assessment Team. 1994. Biodi- 
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Supply and Services. Ottawa. 

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Canadian Council of Forest Ministers (CCFM). 1992. 
Sustainable forests — a Canadian commitment. Minister 
of Supply and Services Canada, Ottawa. 

Fitzsimmons, M, L. Patino, P. MacTavish and P. 
Farrington. 1997. Estimating changes in forest area in 
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Middleton, J. 1994. Effects of forestry on biodiversity in 
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Weyerhauser Canada. 1998. Twenty-year forest manage- 
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Weyerhauser Canada and Saskatchewan Environment and 
Resource Management. Prince Albert, Saskatchewan. 

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Recei\ ed 6 November 2000 
Accepted 19 March 2002 



Status of Redside Dace, Clinostomus elongatus, in the Lynde and 
Pringle Creek Watersheds of Lake Ontario 

Jeff J. Andersen 

Central Lake Ontario Conservation Authority, 100 Whiting Avenue, Oshawa, Ontario LIH 3T3 Canada 

Andersen, Jeff J. 2002. Status of Redside Dace, Clinostomus elongatus, in the Lynde and Pringle creek watersheds of Lake 
Ontario. Canadian Field-Naturalist 1 16(1): 76-80. 

Populations of the Redside Dace, Clinostomus elongatus, are declining throughout most of their Ontario range. Since 
1959, Redside Dace have been found at 20 sampling sites in Lynde and Pringle creeks within the Town of Whitby, 
Ontario. These collection sites were reassessed in 1999 and 2000 to investigate the status of the Redside Dace within 
these systems. A total of 20 Redside Dace were captured at four of the 20 sites. Populations are now either severely limit- 
ed or absent from Pringle Creek and are now confined to upstream areas in Lynde Creek. Redside Dace were confined to 
2"^! and y^ order coolwater streams with variable substrata, and these habitats displayed a 100% canopy closure by grass 
species. Potential Redside Dace habitats are identified within the watershed to provide general guidelines for future land 
development. 

Key Words: Redside Dace, Clinostomus elongatus, fish, minnows, threatened species, special concern, vulnerable species. 



The Redside Dace has a discontinuous global dis- 
tribution and is found only in North America. In the 
western portion of its range, it occurs in the upper 
Mississippi and Lake Michigan drainages, and, in the 
eastern portion, in the Lake Erie, Lake Huron, Lake 
Ontario and upper Susquehanna River drainages. 
Page and Burr (1991) describe the Redside Dace as 
locally common in the eastern part of its range 
although it is declining in many areas. The Redside 
Dace is localized and rare in the western part of its 
range. Collection records for Redside Dace in 
Ontario indicate "the general distribution included 
tributaries of Lake Ontario from Lynde Creek in the 
east to Spencer Creek on the west, and north to the 
headwaters of these systems" and "in the Lake 
Simcoe drainage limited to Kettleby Creek" (Parker 
et al. 1988). They are also known to occur in the 
Saugeen River watershed, Gully Creek and Two Tree 
River in the Lake Huron Drainage, and from Irvine 
Creek in the Lake Erie Drainage. 

The Redside Dace, Clinostomus elongatus, was 
classified as Vulnerable in Canada by the Committee 
on the Status of Endangered Wildlife in Canada 
(COSEWIC) in 1987. In 2000, COSEWIC changed 
the "Vulnerable" classification to Special Concern 
(COSEWIC 2000). Studies are currently underway 
to reassess the status of the Redside Dace on behalf 
of COSEWIC (E. Holm, Royal Ontario Museum, 
personal communications January 1999 to January 
2000). The Ontario counterpart of COSEWIC, the 
Committee on the Status of Species at Risk in 
Ontario (COSSARO) designated the Redside Dace 
as Threatened in Ontario in May 2000 (COSSARO 
2000). 

Redside Dace prefer cool, clear, flowing waters 
with a substrate composed of gravel or stone and are 



reported to be quite sensitive to turbidity (Scott and 
Crossman 1973; Trautman 1981). Redside Dace dif- 
fer from other members of the family Cyprinidae in 
that approximately 77% of their diet consists of 
insects. Daniels and Wisniewski (1994) suggest a 
dominance for flying insects made up mainly of 
danceflies, caddisflies and midges. The Redside 
Dace possesses large eyes and has a very large 
mouth which allows it to catch these in flight by 
leaping out of the water. 

Lynde and Pringle creeks, located in the Township 
of Whitby, Region of Durham, Ontario, centered at 
43° 56' 30" W, 78° 59' 10" N (Figure 1), are the only 
creeks reported to support Redside Dace within the 
Central Lake Ontario Conservation Authority 
(CLOCA) watershed (CLOCA 1996-2000*). 

Redside Dace habitat was described by Parker et 
al. (1988) as pools and slow moving sections of rela- 
tively small headwater streams which have a pool rif- 
fle habitat. Stream sections with overhanging bushes 
and herbaceous plants to provide some cover were 
found to be particularly suitable. Bottom substrates 
were usually composed of boulders, rocks gravel or 
sand, often with a shallow surface covering of detri- 
tus and silt. Streams were clear and colourless in con- 
junction with hard substrates and clear to brown 
tinged streams with organic substrates. This species 
prefers clear water and is quite sensitive to turbidity. 
Novinger and Coon (1998) describe a micro-habitat 
preference for mid-water positions in the deepest 
parts of pools under overhanging vegetation. 

The status of the Redside Dace in Lynde and 
Pringle creeks had not been assessed since 1985. 
Due to the stresses of increased development within 
the Lynde and Pringle creek watersheds and the sen- 
sitivity of Redside Dace to increased turbidity, an 



76 



2002 



Andersen: Redside Dace in Lynde and Pringle Creek Watersheds 



77 



Lynde Creek 
Watershed 




Legend 

- PWentidl Habiuil based nn 
inirared PJiotogr^jby Jnierpretatiori 



J^ ^ ilistortcal and Conlcnqjoraiy Csipturo 

■ - Histork^l Capture 0«ify 
**■• = Watershed Boundaries 
= Roads 



Map Not to Scale 



/ 



Pringle Creek 
Watershed 



Figure 1. Historical (■) and contemporary (*) Redside Dace, Clinostomus elongatus, collection sites and potential habitat 
on Lynde and Pringle creeks. 



impact on the Redside Dace population would 
appear inevitable. The purpose of this study would 
be an attempt to summarize historical capture data, 
resurvey all known capture sites, identify the current 
status and range of known Redside Dace habitat and 
predict potential Redside Dace habitat within the 
Lynde and Pringle creek watersheds. 

Methods 

Historical Capture Mapping 

All historic collection sites from 1959 to 1999 were 
taken from the CLOCA Fisheries Database and 
mapped on a 1:10000 Ontario Base Map (Figure 1). 
Exact locations were difficult to assess due to location 
reporting in previous studies which were given in gen- 
eral or relative terms. Survey locations were, there- 
fore, derived from a combination of the above infor- 
mation, landmarks denoted in field records as well as 
conversations with individuals who conducted the 
studies (E. Holm, personal communication 1999). 

Collection 

Sample collections for the historical studies on the 
Lynde and Pringle were conducted with seine nets 
and dip nets only (Archives of Ontario 1959* and 
Tumey 1984*) or a combination of seine and elec- 
trofishing (Holm and Crossman 1986*), and were 



qualitative in nature. For the current study, qualita- 
tive fish samples were obtained using the single-pass 
electrofishing method described in Stanfield et al. 
(1997). Sampling areas coincided with areas of his- 
toric Redside Dace occurrence. This method was 
effective and least harmful to the fish compared to 
netting methods due to the speed of the sample and 
the relatively short electrical field exposure time. 
The length of steam electrofished ranged from 18 to 
123 metres and averaged 49 metres. The electrofish- 
er was set at 45 Hertz at 4 seconds with an output of 
300 Volts. 

Field collections were conducted July and 
September 1999 and August and October 2000. All 
species captured were noted and Redside Dace cap- 
ture locations were determined using UTM coordi- 
nates. All species were identified in the field and a 
sample of Redside Dace specimens from each cap- 
ture site were sent to the Royal Ontario Museum for 
verification. 

Habitat Mapping 

Potential habitat was identified using Ontario 
Ministry of Natural Resources rectified colour 
infrared photographs (Ontario Ministry of Natural 
Resources 1997). Areas were identified that were 
2nd, 3rd and 4th order streams on the Lynde and 



78 



The Canadian Field-Naturalist 



Vol. 116 



Pringle creeks. These systems were within agricul- 
tural areas with riparian vegetation made up of main- 
ly grass species. 

Results 

Historical Data Summary 

Historically, Redside Dace have been noted in 
both Lynde and Pringle creeks located within the 
Town of Whitby (Figure 1). In four surveys on the 
Lynde and Pringle creeks the capture of the Redside 
Dace was reported (see site codes in Table 1 and 
Figure 1). The first survey was conducted in 1959 by 
the Ontario Department of Planning and Develop- 
ment, wherein five sites on the Lynde Creek and one 
site on Pringle Creek produced Redside Dace (see 
Table 1 site codes preceded by "OPD"). A second 
survey conducted by Tumey in 1983 noted 10 
Redside Dace capture locations on the Lynde (see 
Table 1 site codes preceded by "CLOCA"). The 
third survey produced one confirmed (Royal Ontario 
Museum) Lynde Creek specimen taken by Niblett 
Environmental Associates in 1997 (see Table 1 site 
code preceded by "NBA"). The fourth, and most 
recent study conducted by EcoTec in 1999 noted 
Redside Dace within the Lynde Creek at two sites 
along the Highway #7 corridor (see Table 1, site 
codes preceded by "EcoTec"). Holm and Crossman 
(1985) resurveyed three of the 1959 survey loca- 
tions, and one new site, but no Redside Dace were 
found. They suggested that in Lynde and Pringle 
creeks; (1) there has been a significant decline of C. 
elongatus in the waters sampled, (2) C. elongatus 
has been seriously limited or extirpated from Pringle 
Creek, (3) either the number of individuals, or the 
area of the watershed occupied by the species, has 
declined in Lynde Creek, and (4) there appears to be 
healthy populations in other watercourses. The rec- 
ommendations from this survey concluded; 
"Urbanization definitely seems to reduce the area of 
any watershed which remains suitable to C. elonga- 
tus. It is recommended for streams with known pop- 
ulations of C. elongatus that wherever possible, the 
stream-side vegetation be maintained and instream 
cover and obstructions such as logs not be removed 
(i.e., channelization be avoided), this will maintain 
the pool-riffle habitat and supply a source of terres- 
trial insects — an important food source for the red- 
side dace". 

According to these conclusions and recommen- 
dations, the decline or extirpation of Redside Dace 
in the Lynde and Pringle creek systems appears 
imminent. Lynde and Pringle creeks have been sub- 
ject to many stresses over the years including urban 
development and subsequent channelizations and 
stormsewer outfalls. Many other groups have sur- 
veyed Lynde and Pringle creeks in the vicinity of 
many known Redside Dace occurrences (CLOCA 
1996-2000) and have not captured any specimens 
since 1996, with the exception of the Niblett 



Environmental Associates survey (1997). Redside 
Dace occurrences had not been previously reported 
at the two sites studied by EcoTec (1999*) even 
though the sites had been surveyed prior to 1999. 

Contemporary Study Results 

A total of 20 Redside Dace were collected at four 
of the 20 historical collection sites (Table 1) and 
specimens were verified as C. elongatus (ROM 
Accession #6844). The average total length was 
80.5 mm (ranging between 48-101 mm) and the 
average weight was 4.94g (ranging between 
2-10.1 g) for all samples. At the sites where 
Redside Dace were captured, the following consis- 
tent characteristics of the aquatic and riparian habi- 
tat were noted; 2"'' or 3^^ order streams, a relatively 
steep gradient (1.0-1.59%), riffle pool sequences, 
cool water (summer temperatures between 16.0°C 
and 24.0°C), a streamside vegetation made up of 
grasses providing 100% canopy cover, a lack of 
instream woody debris, a surrounding land use of 
reclaimed agricultural or recreational (golf course) 
and variable substrata including; silt, sand, cobble 
and rock. 

Discussion 

Existing and Potential Habitat 

The four capture sites contained similar habitat. 
It was determined that this habitat type, with some 
exceptions (see below), could be identified in a 
desktop exercise using rectified colour infrared 
aerial photography (Ontario Ministry of Natural 
Resources 1997). Reclaimed agricultural areas that 
were not excessively forested and had riparian veg- 
etation consisting of mainly grass species were 
denoted as potential habitats (Figure 1). The find- 
ings derived from this exercise do not preclude 
other areas and habitat types from being Redside 
Dace terrestrial habitat, rather, they represent areas 
within Lynde and Pringle creeks that contain a 
habitat type that has been proven to support 
Redside Dace. However, there are limitations in the 
"desktop" method of habitat typing: some areas of 
identified potential habitat are either within an 
ephemeral or intermittent watercourse, or do not 
have sufficient flows to support Redside Dace 
(Parker et al. 1988); other areas may have either 
thermal characteristics outside Redside Dace toler- 
ance range, or substrates and riffle/pool characteris- 
tics not normally associated with Redside Dace 
habitat (Scott and Crossman 1973). Furthermore, 
fragmented areas of potential habitat are evident 
within some tributaries and it is unknown if these 
pockets provide enough habitat to be considered 
"potential". For the purpose of this study, these 
areas are included as the potential for Redside Dace 
habitat is evident. It would appear that Redside 
Dace populations are becoming more restricted 
within Lynde Creek and are either severely limited 



2002 



Andersen: Redside Dace in Lynde and Pringle Creek Watersheds 



79 



Table 1. Fish species captured during Redside Dace sampling conducted in Lynde and Pringle creeks in 1999 and 2000. 





















Site Code 
































-o 

c 






















m 


















C3 






















^ 




Species 


150 
CM 

u 

Q 

CU 

O 


•o 
c 
>, 

'^ 

< 

u 

Q 
Cu 
O 


-a 

c 

o 

< 

Z 


1) 
-o 
c 
>. 

J 

vO 
(N 
< 

u 

Q 

CU 

O 


c 

'si- 

< 

U 

o 

u 


■o 
c 
>-. 

r- 

< 

U 
O 

u 


< 

U 
Q 
Cu 
O 

in 
in 

< 

o 

u 


0) 

OS 
u 


'a? 
T3 

c 
>^ 

J 

o 

(N 


c 

< 

U 
Q 

CU 

O 


c 

CO 

OS 
< 

g 

o 


CD 

-o 

c 

oo 

< 

U 
O 

J 

u 


c 

< 


c 
oo 

IT) 

< 

s 

u 


T3 

C 

o 
l/"< 

< 

U 
O 

-] 

u 


c 

< 
U 
O 

u 


C 
< 

U 

o 

u 


c 
oo 

< 

U 

O 

u 


O 
H 


Salmonidae 








































Oncorhynchus mykiss 






18 


4 


1 


1 


4 






4 








4 


16 


1 


2 




55 


Salvelinus fontinalis 








1 












1 










4 




1 




7 


Catostomidae 








































Catostomus commersoni 




9 


10 


17 


22 


9 


4 


7 


8 








7 








2 




95 


Cyprinidae 








































Phoxinus eos 


1 








7 




1 




20 




3 
3 


1 


i 










1 

3 


34 


Clinostomus elongatus 




■%-. 


-* 










13 






■1 


■i 


■ 




20 


Luxilus comutus 








1 


3 


1 


















5 


Pimephales promelas 


11 












4 


5 


20 




1 


3 






2 






2 


48 


Rhinicthys atratulus 


128 


31 


64 


58 


66 


23 


81 


27 


31 


66 


31 


32 


44 


9 


24 


22 


25 


29 


791 


Rhinicthys cataractae 




4 


8 




14 


2 


2 






2 








1 






21 




54 


Semotilus atromaculatus 


17 


85 


63 


60 


46 


24 


37 


82 


52 


4 


28 


29 


58 


8 


5 


1 




60 


659 


Gasterosteidae 








































Culaea inconstans 


3 










1 


2 






















H 


6 


Centrarchidae 




































fir 




Micropterus salmoides 




1 
































K 


1 


Lepomis gibbosus 


2 


1 














3 




3 




11 










V 


20 


Percidae 




































m 




Etheostoma caeruleum 








3 


10 


1 


















2 






m 


16 


Etheostoma nigrum 




10 


14 


22 


98 


20 


8 




2 










3 








K 


177 


Cottidae 




































pi 




Cottus bairdi 




4 


16 


1 


1 






ii 




1 








25 


22 


6 


1 




77 


Total Number 








































of Specimens 


162 


145 


193 167 


268 


82 


143 


134 


136 


78 


69 


65 


121 


50 


75 


30 


52 


95 


2065 


Total Number of Taxa 


6 


8 


7 


9 


10 


9 


9 


5 


7 


6 


6 


4 


5 


6 


7 


4 


5 


5 


16 



or absent within Pringle Creek. Although the range 
of occupation of the Redside Dace within Lynde 
Creek has been reduced, those populations residing 
in suitable habitat appear to be healthy based on 
specimen condition and numbers captured (Table 
1). According to historical field notes, the land use 
surrounding each capture site has remained rela- 
tively unchanged since first being sampled. Those 
sites that have seen a significant land use change 
(i.e., from forested to agricultural or vice-versa) 
since the time of the first sample event did not con- 
tain Redside Dace. It would appear that any 
changes to the above noted habitat characteristics 
would result in a absence of Redside Dace capture. 
One example of this was discovered north of the 
Hamlet of Brooklin wherein a capture site 



described as agricultural in 1959 has been 
reclaimed by an Eastern White Cedar forest by 
1999 and did not display Redside Dace capture. 
Development in and around the Hamlet of Brooklin 
has limited the range of the Redside Dace within 
this area, possibly due to increased sediment load- 
ing and turbidity, riparian vegetation removal and 
temperature changes. 

Population Management Considerations 

Populations of Redside Dace throughout their 
Ontario Range could be subject to a similar exercise 
as reported here to determine the Provincial popula- 
tion status in a consistent manner. Redside Dace 
populations would benefit throughout their range if 
the strictest levels of sediment and turbidity controls 
are utilized during residential and agricultural devcl- 



80 



The Canadian Field-Naturalist 



Vol. 116 



opment. Further areas of study should give consider- 
ation to population estimation, measurements of; 
physical habitat, thermal condition nutrient status, 
dissolved oxygen and turbidity, and the relationship 
of migration barriers to Redside Dace populations. 

Acknowledgments 

It is with gratitude I thank everyone who endeav- 
ored to help me "look for Httle red fish"; Ian Kelsey, 
Owen Kelsey, Mike Wilson, Cyndee Pettifer, Todd 
Langley, Jay Howleg, Erling Holm, Jill McColl, Lori 
Manzon and the Central Lake Ontario Conservation 
Authority. 

Documents Cited [marked * in text] 

Archives of Ontario. 1959. Ontario Department of Plan- 
ning and Development Stream Survey Forms, RGl, HB 
Boxes 2-3. Toronto, Ontario. 

CLOCA (Central Lake Ontario Conservation Author- 
ity). 1996-2000. Stream habitat assessment summaries 
for the CLOCA Watershed. Central Lake Ontario 
Conservation Authority, Oshawa, Ontario. 

EcoTec Environmental Consultants. 1999. Fisheries 
Inventory and Assessment Agreement #2005-A-000017. 
Ministry of Transport, Toronto, Ontario. 

Holm, E., and E. J. Grossman. 1986. A report on a 1985 
attempt to resurvey some areas within the Ontario distri- 
bution of Clinostomus elongatus, the Redside Dace and 
to summarize previous records. Unpublished. Department 
of Ichthyology and Herpetology, Royal Ontario Museum, 
Toronto, Ontario. 

Tumey, P. R. 1984. An inventory and assessment of four 
streams within the Central Lake Ontario Conservation 
Authority. Unpublished. Central Lake Ontario Conser- 
vation Authority, Oshawa, Ontario. 



Literature Cited 

COSEWIC (Committee on the Status of Endangered 
Wildlife in Canada). 2000. Canadian species at risk, 
yearly report. Canadian Wildlife Service, Environment 
Canada, Ottawa, Ontario. 

COSSARO (Committee on the Status of Species at Risk 
in Ontario). 2000. Index list of vulnerable, threatened, 
endangered, extirpated or extinct species of Ontario. 
Queen's Printer for Ontario, Toronto, Ontario. 

Daniels, R. A., and S. J. Wisnievv'ski. 1994. Feeding ecol- 
ogy of Redside Dace Clinostomus elongatus. Ecology of 
Freshwater Fish 1994 3:176-183. 

Novinger, D. C, and Thomas G. Coon. 1998. Behavior 
and physiology of the Redside Dace, Clinostomus elon- 
gatus, a Threatened Species in Michigan. Environmental 
Biology of Fishes. Netherlands 57: 315-326 2000. 

Page, L. M., and B. M. Burr. 1991. A field guide to 
freshwater fishes. Peterson Field Guide Series, Audubon 
Society. Houghton Mifflin Company, New York, New 
York. 432 pages. 

Parker, B. J., P. McKee, and R. R. Campbell. 1988. 
Status of the Redside Dace, Clinostomus elongatus, in 
Canada. Canadian Field Naturahst 102: 163-169. 

Scott, W. B., and E. J. Grossman. 1973. Freshwater fish- 
es of Canada. Bulletin 184, Fisheries Research Board of 
Canada, Ottawa, Ontario. 966 pages. 

Stanfield, L., M. Jones, M. Stoneman, B. Kilgor, J. 
Parish, and G. Wichert. 1997. Stream assessment pro- 
tocol for Ontario. Great Lakes Salmonid Unit, Ministry 
of Natural Resources, Picton, Ontario. 155 pages. 

Trautman, M. G. 1981. The fishes of Ohio with illustrat- 
ed keys. Ohio State University Press, Columbus, Ohio, 
683 pages. 

Received 19 February 2001 
Accepted 22 January 2002 



Aquatic Leaves and Regeneration of Last Year's Straw in the 
Arctic Grass, Arctophila fulva 

Susan G. Aiken' and Rosemary A. Buck- 

'Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario KIP 6P4 Canada; saiken@mus-nature.ca 
-156 Dal ton Trail, Whitehorse, Yukon YIA 3G2 Canada 

Aiken, Susan G., and Rosemary A. Buck. 2002. Aquatic leaves and regeneration of last year's straw in the arctic grass. 
Arctophila fiiWa. Canadian Field-Naturalist 116(1): 81-86. 

Aquatic leaves are reported and described for the grass Arctophila fulva. Re-growth from apparently dead, previous sea- 
son" s straw that had over wintered in the Arctic, is reported. Observations on transplanted material and field observations 
documented that new shoots grew from the apex, and nodes of previous season's stems. Strong underground rhizomes, and 
roots developed resulting in propagules able to develop into new plants. The previous season's straw that regenerates is 
probably detached by animals mid-season, before food reserves in the stems are withdrawn to the rhizomes and roots. Such 
re-generation of detached straw that has over wintered has not been reported previously for any other grass species. 

Key Words: Poaceae, Arctophila fulva, aquatic leaves, vegetative proliferation. Banks Island. 



The grass Arctophila fiilva (Trin.) Rupr. has a low 
Arctic circumpolar distribution (Porsild 1957; 
Tzvelev 1976; Aiken et al. 1996, 2000*). In North 
America, it occurs in Alaska, the Yukon, the 
Northwest Territories, Nunavut, Ontario, Quebec, 
and Labrador. The northern-most record for the 
species in the Western Canadian Arctic Archipelago 
is from Prince Patrick Island, 76°12'N, 119°25'W, 
(Canadian National Herbarium (CAN) 220464). 
Plants grow in shallow, standing water or in wet 
marshy areas where they often form more or less pure 
stands generally 20-50 (and occasionally up to 
100) cm high. The species is easily recognized 
because, unlike any other grasses or sedges that occur 
in the same areas, the mamre plants have leaves that 
are conspicuously larger towards the top of the plant 
than those at the base (Aiken et al. 2000*). 

The occurrence of aquatic leaves in grasses is so 
rare that this is not one of the 1087 characters con- 
sidered for the 9593 taxa in World Grass Species 
(Clayton 1999*). In Canada, wild rice taxa in the 
genus Zizania L. and Signal Grass Pleuropogon 
sabinei R. Br. were the only grasses found to devel- 
op aquatic leaves in the survey done for Watson et 
al. (1986). 

Porsild (1957) noted that A. fulva "near its north- 
ern limit is often sterile, propagating itself vegeta- 
tively". He gave no information on the nature of the 
proliferation and did not describe the propagules. 
Vegetative proliferation by bulbils, formed in place 
of flowers, occurs in the North American arctic 
grasses Poa pratensis (L.) subsp. alpigena (Fr. ex 
Blytt) Hiitonen and Festuca viviparoidea Krajina ex 
Pavlick s.l. (Aiken et al. 2000*). Vegetative prolifer- 
ation by fragmentation of stolons and vertical shoots, 
as a result of goose foraging, appears to be mainly 
responsible for the spread of Puccinellia phrygan- 



odes (Trin.) Scribn. and Merr., a grass not known to 
set seed, and the sedge Carex subspathacea 
Wormsk. that rarely sets seed (Chou et al. 1992; per- 
sonal communication J. Cayouette 2002). 

In these species the establishment of the propag- 
ules occurs in the same growing season. Aerial stem 
fragments can be used as cuttings to produce new 
plants of Bamboos (sample bought in Ottawa 
February 2002), but this is not known for any native 
grasses that occur in Canada, and is unlikely as the 
stems are annual. 

In July 1999, Arctophila fulva was common in 
Aulavik National Park, Banks Island, N.W.T.. 
Canada along the Thomsen River, between 
73°13.49'N, 199°32.12'W and 73°36.2'N, 
120°2' W, as well as at other sites in the park. 

Methods 

At the site on Banks Island, beside the Thomsen 
River (73°46' N, 119°56.7'W), on 10 July 1999, 
plants were growing in a series of shallow pools 
10-70 cm deep surrounded by wet sedge meadows 
predominantly of Eriophorum angustifolium Honck. 
In this pool, most of the new season's growth was 
submerged leaves that if they reached the surface, 
floated parallel to it. Some plants had culms emerg- 
ing from the water and developing short, aerial 
leaves. A sample of each growth form was collected 
and preserved as a herbarium specimen that was 
deposited at the Canadian National Herbarium 
(CAN 582410, Figure 1). 

Beside the tundra pools were pieces of detached 
straw of the previous season's growth that were 
often lying on the damp moss, Hamatocaulis verni- 
cosus (Mitt.) Hideniis. Most of the straw was dead. 
Occasionally, new season's growth, in the form o( 
leafy shoots, was observed at the top of some of the 



81 



82 



The Canadian Field-Naturalist 







Figure \. Herbarium specimen of plants collected N.W.T., 
Banks Island, Aulavik National Park, 73°46' N; 
1 19°56' W, 10 July 1999, S. G. Aiken 99-230 (CAN 
582410). Left, submerged plant with thin, flexuous, 
aquatic leaves and long rhizomes; right, plant devel- 
oping characteristically distichous, more rigid, 
emergent leaves. 



pieces of stranded straw. Samples were collected and 
preserved as a herbarium specimen (CAN 582408, 
Figure 2). Twenty-six additional detached stems 
with new growth, along with some of the moss on 
which stems were lying, were collected. They were 
transported to Banks Island, Sachs Harbour, 
71°59'N, 125°20'W, on 13 July 1999 where the 
straw was sorted into 13 approximately matched 
pairs of stems of similar length, width, and shoot 
development. One set remained in Sachs Harbour 
and the other set was transported to a private home 
in Ottawa-Carleton, Ontario, 45° 16' N, 75°46' W. In 
both locations, the moss was placed in the bottom of 
a shallow container and the stems were laid on top. 
Pond or rainwater was used to saturate the moss to 
duplicate the conditions where the samples were col- 
lected. The containers were placed outdoors in Sachs 
Harbour on 14 July, where temperatures were 
l-3°C, and there was 24-hour daylight and in 
Ottawa on 16 July, where temperatures were 
I7-32°C, and there was a day-night cycle. 
Observations were made as the season progressed. 
When it appeared that lack of nutrients may be limit- 
ing growth, a 10 cm layer of soil was placed under 
the moss on which the culms were lying. 

Results 

Description of aquatic leaves in A.fulva. 

Aquatic or submerged leaves are long, narrow, 
and pale pinkish brown. They look similar to the 
aquatic leaves of the grass Pleuropogon sabinei 



Figure 2. Voucher herbarium specimens collected N.W.T., 
Banks Island, Aulavik National Park, 73°46' N; 
119°56' W, 10 July 1999, S. G. Aiken 99-230c 
(CAN 582408). Left, two typical pieces of previous 
season's straw 25-30 cm long. Right, a stem 
approximately 1 m long, developing new season's 
growth at the apex. Note that the remains of the 
previous season's leaves approximately 15 cm long 
have ends that might be interpreted as chewed. 



R.Br, (image, Aiken et al. 2000*). The leaves were 
suspended in water like those of the aquatic leaves of 
wild rice Zizania palustris (Aiken et al. 1988). 
Submerged leaves have sheaths that are translucent, 
pale pinkish brown, closed to near the apex. Cells 
are developed on 50-60% of the sheath circumfer- 
ence. These are larger in the middle of the tissue and 
small before a thin translucent membrane that forms 
the remaining portion of the circumference. In sur- 
face view cells are readily visible, 2—4.5 mm long, 
0.2-0.3 mm wide, and with prominent end walls. 
Ligules are adnate to the blade for 3-6 mm and taper 
to an acute tip from the blade margins to the mid- 
vein. The free ligule surface is uniformly about 0.5 
mm wide, with the abaxial surface glabrous and the 
margin entire. The ligules are more like those of the 
Cyperaceae than the Poaceae. Collar position is 
inconspicuous. Blades are 8-25 cm long, 0.1-0.4 cm 
wide, translucent, pinkish brown and limp if 
removed from the water. Leaf cells, visible in sur- 
face view, are up to 7 mm long, 0.2-0.3 mm wide 
with prominent end walls. In leaf cross section, the 
midvein is slightly larger than the 8-14 adaxial to 
abaxial girders, 4-7 on either side of the midvein. 
The adaxial surface epidermis is one cell thick; the 
abaxial surface has a single layer of colorless epider- 
mal cells and above them a single layer of cells con- 
taining chlorophyll. Between the epidermi and the 
girders are large air spaces. There is a slight develop- 



2002 



Aiken and Buck: Aquatic Leaves and Regeneration 



83 



ment of cell thickening at both ends of each girder, 
but this is not enough to influence the outline of 
blade surfaces that are uniform in cross section. The 
aquatic leaves observed when the specimens were 
collected have not been described previously in 
North American plants and none was found among 
circumpolar specimens of A. fulva examined by 
Aiken at Kew Herbarium (England, October, 2000). 

Transition and emergent leaves 

These were beginning to form on shoots that had 
started to emerge by 10 July 1999 and had relatively 
stiff leaves less than 10 cm long above the water sur- 
face. Leaves formed near the water surface were 
opaque and greenish brown. Aerial leaves of A. fulva 
have opaque sheaths that are green, brownish green, 
or sometimes reddish, and open to the base. The 
margins are not conspicuously membranous except 
near the sheath apex where the ligule margins are 
decurrent; cells and cell end walls are not visible. 
Ligules are free from the surface of the blade, 2-3 
(-4) mm long slightly higher at the blade margins 
with the abaxial surface minutely scaberulous and 
the apex truncate, lacerate, and ciliolate. The ligule 
is similar to that found in some other arctic Poaceae 
(e.g., Arctagrostis latifolia Aiken et al. 2000*). 
Collars are conspicuous as a zone of contrasting 
colour on the abaxial surface between the sheath and 
the blade. They become more conspicuous in older 
leaves, up to 3 nnm wide at the margins, and narrow 
at the midvein. Blades of the fost aerial leaves are 
3.5^ (-6) cm long and become 10-15 cm long near 
the inflorescence, 0.2-0.8 cm wide, opaque, green, 
spreading stiffly at angles of 45-65° from the culm. 
The position of the vascular bundles is conspicuous 
as ribs on both leaf surfaces, with a lighter colour 
over the ribs, but leaf cells are not visible. In leaf 
cross section, the midvein is often inconspicuous in 
surface view, only slightly larger than the 6-23 
adaxial to abaxial colorless girders. The leaves have 
typical C3 anatomy with zones of chlorenchyma 
between the bundles. Vascular bundles in girders are 
slightly deeper than the chlorenchyma zones 
between the bundles. On top and bottom surfaces of 
the girders are prominent, bubbly zones of compact 
colorless parenchyma. In these positions, the oudine 
of the leaf cross-section has conspicuous contours on 
both surfaces and is 3-4 times as wide as in the 
zones between ribs. 

Last year's dead straw 

Detached stems that had blown to the shoreline of 
the tundra were usually 20-50 cm long, but one mea- 
sured 1 m long (CAN 582408, Figure 2). This stem 
was conspicuously longer than any other grass stems 
seen in Aulavik National Park and may have been 
growing in relatively deep water. There was no evi- 
dence that the A. fulva stems had flowered the previ- 
ous season. Rather, where flowering may have 



occurred the remains of the previous season's leaves 
might be interpreted as having been chewed (Figure 
2). A photograph of the new growth developing at 
the end of the detached stems shows the colour con- 
trast between the previous season's straw colored 
remains and the news season's growth that is both 
reddish and fresh green (image Aiken et al. 2000*). 
On 10 July, none of the nearly 30 sprouting stems 
collected had developed roots. 

Ontario, Ottawa- Carleton. Maximum temperatures 
between 16-23 July 1999, were between 27-32°C. 
By 18 July, one transplanted stem had sprouted a new 
shoot from a stem intemode, and two days later, the 
first root appeared. Five days later, two of the termi- 
nal growing shoots were more than 10 cm high with 
new green growth. By 23 July, there were 24 actively 
growing shoots, the original terminal 13 shoots and 
1 1 new shoots developing at culm nodes. Some were 
less than 1 cm long; the two longest shoots were 
about 15 cm long and curved upwards (Figure 3 A). 
After this, no increase in height occurred in spite of 
warm temperatures in August. At the end of August 
the largest shoots were collected and preserved as 
herbarium specimens. Significant development of 
rhizomes and roots occurred below the moss surface 
during August (Figure 3B) and there was an increase 
in the number of new shoots produced at nodes along 
the previous season's growth until 30 August when a 
total of 5 1 shoots were present. Examination of a lon- 
gitudinal section through apparently dead straw of the 
previous season's growth (Figure 4), indicated the 
stems were brown outside and inside, with little culm 
wall, and hollow on the inside except at the narrow 
node from which the new shoots were developing. 
On 30 August, the largest new stem that had devel- 
oped was harvested and preserved as CAN 582407. 

Banks Island, Sachs Harbour. Maximum tempera- 
tures between 14-24 July 1999, were between 
2.6-13°C. By 24 July, many shoots had begun to 
grow and had red and green leaves. By 24 July, the 
longest shoot and root were 5 cm and 7 cm long, 
respectively. There followed two weeks of sunny 
weather with no rain during which the water in the 
container dried out, the moss turned brown and 
many shoots died. By 1 1 August, there were still 1 3 
shoots alive, some terminal and some developing at 
nodes. The longest shoot was 8 cm and that was the 
maximum height reached. By 23 August, there was 
little sign of more above ground growth and on 3 1 
August, the longest shoot was harvested and pre- 
served as CAN 582556. 

Additional field and herbarium ohserx'ations 

Observations and voucher specimens were col- 
lected in 1999 and 2000 in the course of Parks 
Canada patrols made in Aulavik National Park. 
Figure 5 was photographed on Banks Island, at 
Castel Bay, 74°12'N. 1 19"37'W, on 1 August 1999. 



84 



The Canadian Field-Naturalist 



Vol. 116 




Figure 3. A. Close-up of the terminal end of a leafy shoot approximately 15 cm high that had developed in Ontario by 
1 August 1999, from a vegetatively propagating piece of straw collected on Banks Island. Scale bar in cm. 
B. Close-up of new season's underground growth developing from a node of a previous season's culm (ps). Note 
branching rhizome (rz) with a ring of roots (r) developing from a node (n). 



It shows a vegetative propagule that had developed 
vigorously growing rhizomes and roots. There are 
vouchers for observations made at other three other 
locations in Auluvak National Park, CAN numbers 
582894 - 582896. A table summarizing other field 
observations that were made during Parks Canada 
patrols in 1999 and 2000 has been place with the 
specimens in the A.fulva folder, along with notes doc- 
umenting the occurrence of Muskoxen (Ovibos 
moschatus Zimmerman) and Brant Geese {Branta 
bemicula L.) in the area. In 2001, a field trip in July 
and early August that focused on collecting A. fulva 
and Duponita fisheri R.Br., visited Churchill, 
Manitoba; Coral Habour, Southampton and 
Comwallis Islands, Nunavut; Tuktoyaktuk, and Prince 
Patrick Island, Northwest Territories. At none of these 
places was there any evidence of aquatic leaves or 
vegetative propagules in either grass species. 

Discussion 

Aquatic leaves 

The aquatic leaves in A. fulva observed on Banks 
Island were possibly more obvious because plants 
were growing in water that was relatively deep, not 
much above freezing, and was still cold around 



10 July. Initial growth had been slow and in water 
where the level had not dropped through drying out 
as occurs in shallow ponds on Continental North 
America. The first author observed A. fulva in a 
drained lake-bed on Richard's Island, Northwest 
Territories, 69°20'N, 134°30'W, in 1988, where it 
was the dominant species. The plants were growing 
in damp moss and mud, where there had probably 
not been much depth of water in the spring, tempera- 
tures had been as much as 15°C warmer on average 
than on Banks Island during June, and growth of 
aerial stems appeared to have been rapid. If the rela- 
tively delicate submerged leaves had ever developed, 
they were no longer visible. 

Propagules 

That reserves in detached stems were utilized to 
develop new shoots from living cells that had not 
been killed in the severe winter is a phenomenon not 
known in other Arctic grasses. The vegetative devel- 
opment of the transplanted stems kept on Banks 
Island and those in Ottawa-Carleton, as well as the 
vegetatively propagating specimen found at Castel 
Bay (Figure 5) and other field observations indicate 
that under some environmental conditions A. fulva 
can propagate vegetatively from detached stems. 



2002 



Aiken and Buck: Aquatic Leaves and Regeneration 



85 




Figure 4. A specimen cut longitudinally through the node 
where a new shoot (ns) was developing. Upper por- 
tion with the base of the new leafy stems (ns) was 
arising at the node (n) of an otherwise hollow stem. 
Lower portion is the other side of the stem above. 
The previous season's stem (ps) is hollow, and the 
node (n) is a solid partition across the hollow stem. 



The lack of roots on the sprouting shoots when 
they were collected was probably a reflection of an 
environment where moisture was not limiting 
growth. Early in development, the need of the straw 
of A. fiilva to develop photosynthetic tissue was 
apparently greater than the need to develop roots. 
Although a very different system, the seeds of wild 
rice (Zizania palustris) germinating where water is 
not limiting, also develop photosynthetic tissue 
before producing roots. (Aiken, 1986). The contrast 
in the growth response between the samples 
observed on Banks Island and those observed in 
Ottawa probably reflects the difference in tempera- 
tures at the two locations. The difference in light 
quality or day-length may have been responsible for 
the new growth in Ottawa quickly turning green, 
while the new growth in plants on Banks Island 
remained reddish. In the Ontario transplant, A. fiilva 
behaved as a periodic species (that is, one in which 
development is halted at a particular stage even 
when abundant opportunity for further growth 
remains (Sorenson 1941). It might have been tempt- 
ing to interpret the plants as a genetically dwarfed 
arctic race of the species towards the northern limit 




Figure 5. A piece of previous season's straw successfully 
vegetatively proliferating. It had two rhizomes up to 
12 cm long by 1 August, on Banks Island on the 
coastof Castel Bay, 74° 12' N; 119=37' W. 



of its distribution (Saville 1972) had it not been for 
the previous season's stem 1 m long (Figure 2). 

The detached stems behaved somewhat like 
detached stems of Eurasian Water Milfoil 
(Myriophyllum spicatum L.) that will sometimes 
strand on damp mud. put out roots, and develop 
small terrestrial plants. But in that species, the 
growth occurs in the same growing season (Aiken et 
al. 1979). The specialized over-wintering buds (turi- 
ons) of the native water milfoils {Myriophyllum 
sibiricum Komorov and M. verticillaturm L.). propa- 
gate similarly (Aiken and Waltz 1979; Weber and 
Nooden 1974), but after food reserves have been 
stored in them as they develop. 

Towards northern limits of distribution 

Porsild's (1957) idea that plants of A. fulvo grow- 
ing near the northern limit of distribution of the 
species that had not flowered, may propagate vegeta- 
tively is supported by herbarium vouchers: (a) CAN 
127454 (a collection from Banks Island. 73^24' N, 
1 17° W. A.E. Porsild, 17645, on the label of which 
he noted that the plants were growing in water by a 
shallow pond where perhaps they are always sterile) 
and (b) CAN 127453 (from Victoria Island, near 
70°39' N, 1 17°24' W, made on 8 August 1946, A.E. 
Porsild 17237. on the label of which is noted that the 
sterile specimens were growing in 18" (45 cm) of 
water, forming a margin around a pond and that the 
species is apparently always sterile here). Porsild did 
not entertain the idea that the flowerinu intlores- 



86 



The Canadian Field-Naturalist 



Vol. 116 



cences may have been grazed off and given that the 
culm leaves become longer near the inflorescence the 
top of the culm is not as apparent as in other grasses. 

There are records of A. fulva plants collected at 
similar latitudes that have flowers CAN 5357995 
(Banks Island, Egg River 72°27' N, 124°36'), CAN 
535769 (Banks Island, Shoron Lake, 73°N, 
124°18'W) and CAN 203530 (the northernmost 
specimen from Prince Patrick Island 76°12'N). 

Animal grazing 

Arctophila fulva is utilized as a forage by 
Muskoxen and Brant Geese (CAN 220464 and Park 
Canada patrol observations in Aulavik National 
Park). It is possible that plants had begun to flower, 
or were flowering, but. the inflorescences were 
removed by some of these animals, grazing. Gray 
(1987) described Muskoxen playing in water on 
Bathurst Island (where A. fulva does not occur). He 
had observed that animals would walk into the mid- 
dle of a shallow pond, stand in the water and sudden- 
ly begin to head-toss, whirl around, jump and splash. 
If A. fulva was growing in such a pond, plants would 
likely be uprooted by such activities. The tundra 
pools observed on Banks Island would be ideal for 
such play and the large Muskoxen population on the 
island may well explain the large number of 
detached stems found around the margins of ponds. 
These may float to the surface, and later be blown to 
the shoreline of the ponds, where they over-winter. 
Possibly some stems are detached when they are 
actively growing and contain food reserves that oth- 
erwise, later in the season, would have been with- 
drawn to the rhizomes and roots with the onset of 
winter. Such reserves appear to have been available 
to initiate new growth the following season, in some 
detached stems. On both Banks and Victoria islands, 
Muskoxen and Brant Geese are common and their 
activities may contribute to A. fulva occasionally 
proliferating vegetatively from detached straw. 
When the phenomenon of straw from one season 
being able to sprout new plants in the next season 
was discussed with grass taxonomists at Kew, they 
were unaware of any other such occurrence and 
thought it most unlikely to happen in tropical or tem- 
perate areas (W. D. Clayton, S. M. Phillips, and S. 
A. Renoize, personal communication March 2002). 

Acknowledgments 

The authors acknowledge with thanks the logistic 
support of this project by Parks Canada, and particu- 
larly by N. A. Lawrence, the Chief Warden of 
Aulavik National Park, 1999-2000, Polar 
Continental Shelf Project, and the Canadian Museum 
of Nature. They also acknowledge permission given 
to carry out plant research in the Park by Parks 
Canada, the Hunters and Trappers Committee, Sachs 
Harbour, and the Environmental Impact Screening 
Committee, Inuvik. The research was undertaken on 
the science license #13055N from the Aurora 



Research Institute. For help with producing figures, 
thanks go to R. Boles and J. Madill. Anonymous 
reviewers and W. D. Clayton, S. M. Phillips, and 
S. A. Renoize are thanked for helpful discussions on 
earlier versions of the manuscript. 

Documents Cited (Marked * in text) 

Aiken, S. G., L. L.Consaul, and M. J. Dallwitz. 2000 
onwards. Grasses of the Canadian Arctic Archipelago: 
Descriptions, Illustrations, Identification and Information 
Retrieval. URL [RTF bookmark start: _Hlt492 185404] 
[RTF bookmark end: _Hlt492 1 85404]/delta/arcticf . Hard 
copy deposited at the Canadian Museum of Nature 
Library. 

Clayton, W. D. 1999. World Grasses Database v. 5.1 
http://www.rbgkew.org.uk/data/grasses. 

Literature Cited 

Aiken, S. G. 1986. The distinct morphology and germi- 
nation of the grains of two species of wild rice (Zizania, 
Poaceae). Canadian Field-Naturalist 100: 237-240. 

Aiken, S. G., L. L. Consaul, and M. J. Dallwitz. 1996. 
Grasses of the Canadian Arctic Archipelago: a DELTA 
database for interactive identification and illustrated 
information retrieval. Canadian Journal of Botany 74: 
1812-1825. 

Aiken, S. G., P. F. Lee, D. Punter, and J. M. Stewart. 
1988. Wild rice in Canada. NC Press Ltd. Toronto. 130 
pages. 

Aiken, S. G., P. R. Newroth, and L Wile. 1979. Biology 
of Canadian Weeds. 34. Myriophyllum spicatum L. 
Canadian Journal of Plant Science 59: 201-215. 

Aiken, S. G., and K. F. Waltz. 1979. Turions of 
Myriophyllum exalbescens Fernald. Aquatic Botany 6: 
357-363. 

Chou, R., C. Vardy, and R. L. Jefferies. 1992. Establish- 
ment from leaves and other plant fragments produced by 
the foraging activities of geese. Functional Ecology 6: 
297-301. 

Gray, D. R. 1987. The Muskoxen of Polar Bear Pass. Co- 
published by the National Museum of Natural Sciences, 
National Museums of Canada, Ottawa, and Fitzhenry & 
Whiteside, Markham, Ontario 191 pages. 

Porsild, A. E. 1957. Illustrated flora of the Canadian Arctic 
Archipelago. National Museum of Canada Bulletin 
Number 146. 

Saville, D. B. O. 1972. Arctic adaptations in plants. Re- 
search Branch, Canada Department of Agriculture, 
Monograph 6. 

S0rensen, T. 1941. Temperature relationships and phe- 
nology of the Northeast Greenland flowering plants. 
Meddelelser om Gr0nland 125: 1-305. 

Tzvelev, N. N. 1976. Grasses of the Soviet Union [Zlaki 
SSSR. In Russian.] Nauka, Leningrad. 

Watson, L., S. G. Aiken, M. J. Dallwitz, L. P. Lefkovitch 
and M. Dube. 1986. Canadian grass genera: keys and 
descriptions in English and French from an automated 
data bank. Canadian Journal of Botany 64: 53-70 + 2 
microfiche Data Banks approx. 900 pages. 

Weber, J. A. and L. D. Nooden. 1974. Turion formation 
and germination in Myriophyllum verticillatum; phenolo- 
gy and its interpretation. Michigan Botanist 13: 151-158. 

Received 20 February 2001 
Accepted 14 March 2002 



Records of Northern Mockingbird, Mimus polyglottos. 
Occurrences in North Dakota During the Twentieth Century 

Lawrence D. Igl* and Ron E. Martin^ 

'Northern Prairie Wildlife Research Center, U.S. Geological Survey, 871 1 37th Street SE, Jamestown, North Dakota 58401 

USA 
216900 125th Street SE, Sawyer, North Dakota 58781 USA 

Igl, Lawrence D., and Ron E. Martin. 2002. Records of Northern Mockingbird, Mimus polyglottos, occurrences in North 
Dakota during the twentieth century. Canadian Field-Naturalist 1 16(1): SI -91. 

The Northern Mockingbird {Mimus polyglottos) is a common bird in the southern United States that has been expanding its 
breeding range into the northern United States and southern Canada. During the twentieth century, there were 128 reports 
of Northern Mockingbird occurrences in North Dakota, including 106 reports during the breeding season (15 April to 31 
August) and 22 during the nonbreeding season (1 September to 14 April). The species has been largely absent from North 
Dakota from January through mid- April. Prior to the 1930s, there was only one record (1916) of the Northern Mockingbird 
in the state. Observations of Northern Mockingbirds in North Dakota increased markedly between the 1930s and 1990s. On 
average, there were 0.3 reports of mockingbirds per year in 1931-1940. 0.6 in 1941-1950, 1.1 in 1951-1960, 1.6 in 1961- 
1970, 2.4 in 1971-1980, 2.3 in 1981-1990, and 4.5 in 1991-2000. The species has been observed in North Dakota nearly 
annually since 1958. At least six reports during the twentieth century included evidence of nesting (nests or dependent 
young). Based on mockingbird records during the twentieth century, we designate the current status of the Northern 
Mockingbird in North Dakota as a rare spring migrant, rare summer visitant, casual nester, and a casual fall and winter 
visitant. 

Key Words: Northern Mockingbird, Mimus polyglottos, distribution and abundance. North Dakota, population status, 
twentieth century. 



The Northern Mockingbird {Mimus polyglottos) is 
a geographically widespread species with southern 
affinities in North America. Densities of the species 
are highest near the Gulf of Mexico and decline 
northward and westward (Price et al. 1995). During 
the twentieth century, the species expanded the 
northern limits of its range into the northern United 
States and southern Canada (Derrickson and 
Breitwisch 1992). Although mockingbirds are con- 
sidered permanent residents throughout much of 
their historical range, mockingbirds are believed to 
be migratory in the northern portion of their range, 
including newly colonized northern regions (Brazier 
1964; Stiles 1982; David et al. 1990; Derrickson and 
Breitwisch 1992). 

Stevens (1947, 1948) summarized some of the 
earliest records of Northern Mockingbirds in North 
Dakota. Brazier (1964) reviewed the range expan- 
sion of the Northern Mockingbird in the northern 
Great Plains and provided updated and new informa- 
tion on the Northern Mockingbird in North Dakota. 
Stewart (1975) summarized historical occurrences of 
the species and categorized the Northern 
Mockingbird as a hypothetical breeder in the state. 
Little has been published on mockingbirds in North 
Dakota since Stewart's book. There has been a sub- 
stantial increase in the number of reports of Northern 
Mockingbirds in North Dakota (e.g., Johnson and 
Johnson 1976), including confirmed nesting records 
(e.g., Child et al. 1980). In this paper, we summarize 



reported occurrences of Northern Mockingbirds in 
North Dakota during the twentieth century, review 
the species' current status and distribution within the 
state, and discuss factors influencing or limiting the 
species' expansion into the state. 

Methods 

Records of Northern Mockingbird occurrences in 
North Dakota were summarized from published and 
unpublished reports, including those in Bird Lore, 
Audubon Magazine, Audubon Field Notes, American 
Birds, and National Audubon Society Field Notes, 
and those filed with the North Dakota Birding 
Society (R. E. Martin and G. B. Berkey, North 
Dakota Birding Society, unpublished data). We also 
included data from Christmas Bird Counts (1909- 
1999; http://birdsource.cornell.edu/cbc/). North 
American Breeding Bird Surveys (1967-1999; Sauer 
et al. 2000; U.S. Geological Survey. Laurel, 
Maryland, unpublished data), and banding records 
from the Bird Banding Laboratory (1955-1998; U.S. 
Geological Survey. Laurel. Maryland, unpublished 
data). We defined a record as an occurrence of an 
individual, a breeding pair, or a group of Northern 
Mockingbirds. Records were summarized by date 
(decade and month) and location (county) of the 
observation. Details on nests or dependent young 
also were noted. Exact locations or dales are not 
known for a few records. Although some observa- 
tions occurred over multiple days, weeks, or months. 



87 



88 



c 
o 



CO 

O 



50 



40- 



<D 30-1 



20- 



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Vol. 116 



B Breeding Season 
I I Nonbreeding Season 




1901 1911 1921 1931 1941 1951 1961 1971 1981 1991 

Decade 



Figure 1 . Records of Northern Mockingbird occurrences in North Dakota during the 
eth century, by decade (e.g., 1901 to 1910, 1991 to 2000). Dark = breeding 
records; light = nonbreeding-season records. 



twenti- 
-season 



summaries within figures represent the date that the 
individual record was first observed. For purposes of 
discussion, observations between 15 April and 31 
August were considered breeding-season records, 
and observations between 1 September and 14 April 
were considered nonbreeding-season records 
(Derrickson and Breitwisch 1992) . 



Results 

During the twentieth century, there were 128 
reports of Northern Mockingbirds in North Dakota 
(Appendix A). The number of individuals detected 
during a single observation ranged from one to four. 
Most of the observations involved single birds 
observed for only one or a few days. 




M J J A 

Month 



O N D 



Figure 2. Records of Northern Mockingbird occurrences in North Dakota during the 
twentieth century, by month. 



I 



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Igl and Martin: Mockingbird Occurrences in North Dakota 



89 




Figure 3. Northern Mockingbird nest found on 25 June 1998 in Kidder County, North Dakota (photograph by L. D. Igl). 



The earliest confirmed record of a Northern 
Mockingbird in North Dakota was of an adult bird 
collected on 23 November 1916 by C. C. Smith on 
the campus of the University of North Dakota in 
Grand Forks (Grand Forks County) (Wood 1923). 
Wood (1923: 77) considered this bird to be an "acci- 
dental straggler." To our knowledge, there were no 
records of Northern Mockingbirds in North Dakota 
before that date. This observation also represented 
the first record of the species in the northern Great 
Plains during the nonbreeding season (Brazier 1964). 

The second record of a Northern Mockingbird in 
North Dakota occurred over 17 years later on 17 
May 1934 in Valley City (Barnes County) 
(Appendix A). This observation represented the first 
breeding-season record in the state and generally 
marked the beginning of the species' range expan- 
sion into North Dakota. Observations of mocking- 
birds increased markedly between the 1930s and 
1990s (Figure 1). On average, there were 0.3 reports 
of mockingbirds per year in 1931-1940, 0.6 in 1941- 
1950, 1.1 in 1951-1960, 1.6 in 1961-1970, 2.4 in 
1971-1980, 2.3 in 1981-1990, and 4.5 in 1991-2000. 
Since 1958, the species has been observed nearly 
annually in all but 3 of 43 years, with no records for 
1962, 1983, and 1989. 

Only 22 mockingbird occurrences were recorded 
during the nonbreeding season, mostly between 



October and December (Figure 2: Appendix A). 
Mockingbirds were largely absent from the state in 
January, February, and March. Only two mocking- 
birds have been observed in North Dakota during 
those months. Between 30 November 1972 and 14 
April 1973, an injured mockingbird was observed in 
Fargo in Cass County. Between mid-December 1999 
and 1 February 2000, a single mockingbird sporadi- 
cally visited bird feeders about 6.4 km southeast of 
Jamestown in Stutsman County (M.A. Sovada, U.S. 
Geological Survey. Jamestown, North Dakota, per- 
sonal communication). 

About 83% (106 of 128) of the mockingbird obser- 
vations occurred during the breeding season (mid- 
April through August) (Figure 2, Appendix A). Of 
the 106 reports of mockingbird occurrences in North 
Dakota during the breeding season, six (6%) involved 
nests or dependent young. Mrs. R. Nordbye (personal 
communication in Child et al. 1980) found a nest 
with young in Parshall (McLean County) in the sum- 
mer of 1961. Near Richardton (Stark County). J. Hoff 
found nesting mockingbirds in July 1970. June 1988. 
and July 1995. Successful fledging of two successive 
broods in June and Julv 1980 was reported in Oakes 
in Dickey County (Childs et al. 1980). L. D. Igl 
found a breeding pair and a nest containing four eggs 
in a single-row shelterbell in Kidder County on 25 
June 1998 (Figure 3; Martin 1998). There also are 



90 



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The Canadian Field-Naturalist 



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30 60 km 



Figure 4. Records of Northern Mockingbird occurrences in North Dakota during the twentieth 
century, by county. Asterisk (*) denotes counties where nests or dependent young 
have been observed. 



two references to mockingbirds nesting in North 
Dakota but without specific reference to date: A. C. 
Fox indicated that mockingbirds have been found 
nesting in Dickinson (Stark County; Stewart 1975), 
and R. L. Rytter noted adult mockingbirds feeding 
young in Kenmare (Ward County). 

Northern Mockingbirds were observed in 36 
(67.9%) of the 53 counties in North Dakota (Figure 
4, Appendix A). Although more mockingbird obser- 
vations occurred in the southern half than the north- 
ern half of North Dakota (Figure 4), the temporal 
pattern of expansion into North Dakota was sparse 
and sporadic rather than progressing from south to 
north (Appendix A). Counties with larger cities had 
more records of mockingbirds than less-populated 
counties. These included Bismarck in Burleigh 
County, Mandan in Morton County, Fargo in Cass 
County, Jamestown in Stutsman County, Minot in 
Ward County, Dickinson in Stark County, and Grand 
Forks in Grand Forks County. The largest number of 
records (n = 26) in a county occurred in Cass 
County, with most occurrences being from Fargo, 
the largest metropolitan area in North Dakota. The 
distribution of mockingbird records within the state 
also may reflect the distribution of observers in 
North Dakota. For example, there were seven 
records of mockingbirds in Adams County in south- 
western North Dakota, all made by C. Griffiths 
and/or D. Griffiths. 

The North American Breeding Bird Survey (BBS) 
is a program of standardized roadside surveys 



administered by the U.S. Geological Survey and 
conducted by experienced or knowledgeable volun- 
teers, usually in June. BBSs have been conducted in 
North Dakota since 1967; there are currently 44 
active routes in the state. Northern Mockingbirds 
were recorded on 5 of 983 (< 1%) surveys of BBS 
routes that were conducted in North Dakota between 
1967 and 1999, including one year each on the 
Parshall (McLean County), Denbigh (McHenry 
County), and Sheyenne Lake (Sheridan County) 
routes and two years on the Bentley (Hettinger 
County) route (Appendix A). The first occurrence of 
a mockingbird on a BBS route was on the Parshall 
route on 19 June 1985. 

Sponsored by the National Audubon Society, 
Christmas Bird Counts (CBCs) are conducted annu- 
ally throughout North America. The main goal of the 
CBC program is to monitor bird populations during 
early winter (late December to early January). CBC 
circles are frequently placed where the greatest con- 
centrations of wintering birds occur and often are 
placed near towns and cities. CBCs have been con- 
ducted in North Dakota since the early part of the 
twentieth century. The first CBC in North Dakota 
occurred in Fargo in 1909 (Chapman 1910); one or 
more CBCs have been conducted nearly annually in 
North Dakota since then. Mockingbirds have been 
reported on 4 of 682 (< 1%) CBCs in North Dakota 
between 1909 and 1999, including three years on the 
Bismarck-Mandan CBC and one year on the Fargo- 
Moorhead CBC (Appendix A). Mockingbirds also 



2002 



Igl and Martin: Mockingbird Occurrences in North Dakota 



91 



were reported within the Count circle during the 
week of the CBC but not the day of the CBC in 
Fargo (Cass County) in 1972 and in Jamestown 
(Stutsman County) in 1999. The earliest occurrence 
of the species on a CBC was in Bismarck (Burleigh 
County) on 20 December 1956. 

To our knowledge, only four Northern 
Mockingbirds have been banded in North Dakota. A 
mockingbird was banded by H. R. Gray in Wilton 
(McLean County) on 3 June 1943 (Stevens 1948). 
Of 610 066 birds banded in North Dakota between 
1955 and 1998 (all species combined), three were 
Northern Mockingbirds banded in 1960, 1964, and 
1971 by R. T. Gammell and A. M. Gammell (Bird 
Banding Laboratory, U.S. Geological Survey, 
Laurel, Maryland, unpublished data). 

Discussion 

The available data suggest that North Dakota cur- 
rently supports a small but growing population of 
Northern Mockingbirds. Spring arrival in North 
Dakota begins in mid- April and continues into May 
(Figure 2), and available evidence suggests that nest- 
ing occurs largely in June and July. One or two 
broods are possible, but rare, in North Dakota. Few 
mockingbirds have overwintered in the state, which, 
along with the influx of mockingbirds in spring, sug- 
gests that this species is migratory in North Dakota. 
However, most of the North Dakota observations 
were of single birds, and to our knowledge, most 
birds were observed or present for only one or a few 
days. Observations of mostly single birds and the 
general scarcity of nesting suggests that most mock- 
ingbirds observed in North Dakota were probably 
wandering rather than nesting birds. Mockingbirds 
are noted for their behavioral plasticity and wander- 
ing behavior (Derrickson and Breitwisch 1992). 

Many records of mockingbird occurrences in 
North Dakota were from scattered or random obser- 
vations by one or a few observers. Most observations 
occurred in spring or summer, and many observa- 
tions occurred in or near human population centers. 
Admittedly, these records may be biased by the 
activity and locations of observers (i.e., most birding 
occurs in spring and summer and most bird observa- 
tions are made in residential areas). Apparent 
increases in North Dakota during the twentieth cen- 
tury also may reflect higher observer effort or more 
qualified observers. However, mockingbirds are a 
conspicuous and easily detected species. 
Mockingbirds strongly defend their territories during 
both the nonbreeding and breeding seasons, vocalize 
loudly and elaborately, and often are associated with 
human residential areas. Thus, increases in mocking- 
bird populations based on individual records proba- 
bly track statewide trends. 

Changes in mockingbird populations in North 
Dakota also coincide with similar patterns of range 



expansion in other states and Canadian provinces. 
During the twentieth century, the species has 
expanded its range northward in California (Arnold 
1980), Michigan (Dziepak 1991), New York (Bull 
1974), Ontario (Curry 1987), Quebec (David et al. 
1990), Saskatchewan (Smith 1996), and Wisconsin 
(Robbins 1991). There are scattered records of 
mockingbirds in states adjacent to North Dakota, 
including Montana (MBDC 1996), South Dakota 
(SDOU 1991), and Minnesota (Janssen 1987). In 
western Illinois, the species was historically an irreg- 
ular migrant until the mid- 1930s, when it became 
common (Bohlen 1989). Mockingbirds were sparse 
in Iowa until the 1930s (Spess Jackson et al. 1996); 
the species is now considered a rare summer resident 
with higher densities in the southern portion of the 
state (Kent and Dinsmore 1996). 

Factors thought to contribute to the species' north- 
ward range expansion in North America include 
increased availability of suitable nesting habitat 
(trees and shrubs) and increased availability of food 
(e.g., fruit-bearing trees and shrubs, suet feeders). 
Northern Mockingbirds favor brushy areas and 
woodland edges, and often are associated with orna- 
mental plantings in residential areas (Derrickson and 
Breitwisch 1992) and shelterbelts and scattered trees 
and shrubs in agricultural areas (Versaw 1998). 

The abundance and distribution of woody vegeta- 
tion have changed dramatically in North Dakota 
since this region was first settled by Europeans in the 
mid to late 1800s. Since settlement, there have been 
repeated efforts to establish tree plantings in North 
Dakota and other areas in the Great Plains (Hart and 
Hart 1997). Fire suppression also encouraged the 
encroachment and establishment of woody vegeta- 
tion into open grasslands in this region. Between 
1935 and 1942, 38 million trees were planted in 13 
760 ha of shelterbelts and windbreaks (Haugen 
1999). Since the Dust Bowl, more than 142 000 km 
of field windbreaks and 1 1 5 000 ha of protection tree 
plantings have been established in North Dakota. It 
is probable that the planting of shelterbelts and wind- 
breaks in North Dakota in the early part of the twen- 
tieth century provided the habitat necessary and 
impetus for the Northern Mockingbird's northward 
expansion into North Dakota. Presently, there does 
not appear to be a shortage of suitable breeding habi- 
tat for Northern Mockingbirds in North Dakota. 

Fruit is a critical component of the mockingbird's 
diet during the fall, winter, and spring (Derrickson 
and Breitwisch 1992). Stiles (1982) speculated that 
the species' northward expansion into the northeast- 
ern U.S. may have been related to the plantings of 
multiflora rose {Rosa niultiflora). In the northern 
Great Plains, mockingbirds have been observed 
feeding on fruit from several common fruit-bearing 
trees and shrubs including honeysuckle {Loniceni 
spp.), Russian Olive {EUwci^nus ungustijoliu), and 



92 



The Canadian Field-Naturalist 



Vol. 116 



apple (Malus spp.) (Brazier 1964). In other areas, the 
species has been observed eating fruit from 
hawthorn {Crataegus spp.), crabapples {Pyrus spp.), 
and Eastern Redcedar {Juniperiis virginiana) 
(Arnold 1980; Stiles 1982), all of which occur in 
North Dakota. Dziepak (1991) indicated that suet 
feeders and ornamental plantings of fleshy-fruited 
shrubs may have helped the species expand its range 
into Michigan, but these food resources were not 
sufficient to sustain the species through Michigan's 
severe winters. 

Northern Mockingbirds are largely absent from 
North Dakota during the winter. Overwintering also 
is rare in other northern regions (e.g., Saskatchewan; 
Smith 1996), suggesting that mockingbirds withdraw 
from these areas for the winter or succumb to winter 
conditions. Dziepak (1991) and others have suggest- 
ed that winter weather is likely the chief factor limit- 
ing the distribution of this species at the northern 
edge of its range. Root (1988) proposed that a physi- 
ological limit on metabolic rate constrains the north- 
ern limit of wintering North American birds, and 
demonstrated that mockingbirds typically overwinter 
in areas that have an average minimum January tem- 
perature of -7°C or higher. The average January 
temperature in North Dakota ranges from -16.7°C in 
northeast North Dakota to -8.3°C in southwest 
North Dakota (Jensen 1972). Repasky (1991), how- 
ever, argued that the northern boundary of a species 
would occur at the point where sufficient food can- 
not be obtained to offset the greater energy demand 
of colder temperatures. Brazier (1964) noted many 
records of dead mockingbirds found during winter in 
the northern Great Plains. In Illinois, mockingbirds 
have suffered massive reductions in population dur- 
ing severe winter weather (Bohlen 1989). The suc- 
cess of the species' range expansion into northern 
areas will likely be determined by the species' ability 
to develop stronger migratory habits or to cope with 
the harsh winter conditions and limited food 
resources in northern regions. 

Based on mockingbird records of occurrence in 
North Dakota during the twentieth century, we des- 
ignate the current status of the Northern Mock- 
ingbird in North Dakota as: (1) a rare spring migrant 
(i.e., a species that occurs annually in North Dakota 
in low numbers during spring), (2) a rare summer 
visitant (i.e., a species that wanders or oversummers 
annually in low numbers in North Dakota during the 
normal breeding period), and (3) a casual fall and 
winter visitant (i.e., a species that occurs less than 
annually in North Dakota during fall or winter). 
Although there were very few nesting records in 
North Dakota during the twentieth century, we also 
recommend that the species' nesting status in North 
Dakota be changed from "hypothetical breeder" 
(Stewart 1975) to "casual nester" (i.e., a species for 
which a viable clutch of eggs, dependent young in 
the nest, or dependent young that have left the nest 



have been observed less than annually in North 
Dakota). Continued monitoring of mockingbirds 
should provide insight into the dynamics of the 
species' range expansion into North Dakota and 
elsewhere in the northern Great Plains. Future 
increases or changes in the species' distribution and 
abundance in North Dakota will necessitate a re- 
evaluation of the species' status. 

Acknowledgments 

Over the years, many people have contributed 
records of Northern Mockingbirds in North Dakota. 
We thank all of them. We are especially grateful to 
Gordon B. Berkey and David O. Lambeth for their 
dedication to North Dakota birdlife, for their com- 
mitment to the North Dakota Birding Society, and 
for compiling bird records for the state from the mid- 
1970s to the mid-1990s. This paper has benefitted 
from the thoughtful comments of G. B. Berkey, F. R. 
Cook, A. J. Erskine, C. S. Houston, D. H. Johnson, 
H. T. Sklebar, and M. A. Sovada. 

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Arnold, J. R. 1980. Distribution of the mockingbird in 

California. Western Birds 11: 97-102. 
Berkey, G. B. 1982. Northern Great Plains. American 

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Berkey, G. B. 1984. Northern Great Plains. American 

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4-11. 
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Research paper NC-336. 101 pages. 
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94 



The Canadian Field-Naturalist 



Vol. 116 



Versaw, A. E. 1998. Northern Mockingbird (Mjm«5 /7o/y- 
glottos). Pages 400-401 in Colorado breeding bird atlas. 
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Received 21 March 2001 
Accepted 22 March 2002 



Appendix A. 

Location, date(s), observer(s), season (nonbreeding or breeding), and number of birds observed for reported occurrences of 
Northern Mockingbirds in North Dakota during the twentieth century. Data were summarized from published and unpub- 
lished (R. E. Martin and G. B. Berkey, North Dakota Birding Society) reports. 



No. 


Location 


Date(s) 


Observer(s) 


Season^ 


Number of birds 


1. 


Grand Forks, Grand Forks County 


23 November 1916 


C. C. Schmidt (Woods 1923) 


Nonbreeding 


one (collected) 


2. 


Valley City, Barnes County 


17 May 1934 


Observer not specified 
(from the files of the 


Breeding 


one 








North Dakota Birding Society) 




3. 


Dickinson, Stark County 


Summer 1938 


R. W. Smith (Stevens 1947) 


Breeding 


one 


4. 


Dickinson, Stark County 


May 1939 


R. W. Smith, F. C. Butcher, 
and F. G. Butcher 
(Stevens 1947) 


Breeding 


one 


5. 


Fargo, Cass County 


12-13 May. 1942 


Mrs. W. E. Brentzel 


Breeding 


one 


6. 


Rock Lake, Towner County 


31 May 1943 


Ambrosen and Dinkins 
(Hammond 1943) 


Breeding 


one 


7. 


Minot, Ward County 


1-2 June 1943 


E. A. Hibbard (Stewart 1975) 


Breeding 


one 


8. 


Wilton. McLean County 


3 June 1943 


H. R. Gray (Stevens 1948) 


Breeding 


one (banded) 


9. 


Fargo, Cass County 


21-28 July 1946 


0. A. Stevens and 
H. G. Heggeness 
(Stevens 1947) 


Breeding 


one 


10. 


Minot, Ward County 


1-2 June 1948 


S. Saugstad (Stewart 1975) 


Breeding 


one 


11. 


Mandan, Morton County 


24-25 May 1952 


R. N. Randall 


Breeding 


one 


12. 


Kenmare, Ward County 


1-14 June 1952 


R. T. Gammell and 

A. M. Gammell 

(Gammell and Gammell 1952) 


Breeding 


one 


13. 


Bismarck, Burleigh County 


8 June 1952 


R. N. Randall 
(Baumgartner 1952) 


Breeding 


one 


14. 


Fargo, Cass County 


13 May 1953 


J. F. Cassel (Gammell and 
Gammell 1953) 


Breeding 


one 


15. 


Des Lacs National Wildlife 
Refuge, Ward County 


4-7 October 1955 


H. S. Huenecke (Gammell 
andHuenecke 1956) 


Nonbreeding 


one 


16. 


Southeast of Kenmare, 
Ward County 


18 May 1956 


V. Rytter (Gammell 1956) 


Breeding 


one 


17. 


Mandan, Morton County 


30 December 1956 


Bismarck-Mandan Christmas 


Nonbreeding 


one 








Bird Count (Cruickshank 1957) 




18. 


Parshall, McLean County 


4-1 1 November 1958 Dr. and Mrs. R. Nordbye; 


Nonbreeding 


one 








R. T. Gammell and 












A. M. Gammell (Krause 1959) 






19. 


Des Lacs National Wildlife 
Refuge, Ward County 


23 August 1959 


A. M. Gammell 
(Krause 1960) 


Breeding 


four 


20. 


Ward County 


6 May 1960 


R. T. Gammell 


Breeding 


one (banded) 


21. 


Kenmare, Ward County 


25-28 May I960 


R. L. Rytter and D. Rytter 
(Gammell I960) 


B receding 


one each at 
three locations 


22. 


Parshall, McLean County 


Summer 1961 


Mrs. R. Nordbye (personal 
communication //; Child et al. 
1980) 


Breeding 


nest with young 


23. 


Mandan, Morton County 


29 December 1963 


Bismarck-Mandan 
Christmas Bird Count 
(Cruickshank 1964) 


Nonbreeding 


one 


24. 


Ward County 


1 May 1964 


R. T. Gammell 


Breeding 


one (banded) 


25. 


Bismarck. Burleigh County 


10 May 1964 


R. N. Randall 


Breeding 


one ^^ 


26. 


Fargo, Cass County 


13 June 1965 


Mrs. F. B. Scheel 
(Hatch 1965) 


Breeding 


one ^^ 


27. 


Near Binford. Griggs County 


June 1966 


R. E. Stewart, Sr. 
(Stewart 1975) 


Breeding 


one 












(Continued) 



2002 



Igl and Martin: Mockingbird Occurrences in North Dakota 



95 



Appendix A. (continued) 



No. 


Location 


Date(s) 


Observer(s) 


Season^ 


Number of birds 


28. 


Hankinson, Richland County 
(T130N,R50W) 


12 July 1966 


R. E. Stewart, Sr. 
(Stewart 1975) 


Breeding 


one 


29. 


North of Fargo, Cass County 


30 April- 19 

May 1967 


R. Kroodsma 


Breeding 


one 


30. 


Bismarck, Burleigh County 


20 May 1967 


R. N. Randall 


Breeding 


one 


31. 


Fargo, Cass County 


22-23 May 1967 


Observer not specified 

(from the files of the 

North Dakota Birding Society) 


Breeding 


one 


32. 


Rhame, Bowman County 


20 June 1967 


R. E. Stewart, Sr. 
(Stewart 1975) 


Breeding 


one 


33. 


Fargo, Cass County 


13 September 1967 


Observer not specified 

(from the files of the 

North Dakota Birding Society) 


Nonbreeding 


one 


34. 


Arrowwood National 

Wildlife Refuge, Stutsman County 


21 April 1968 


J. T. Lokemoen and 
P. F. Springer 


Breeding 


one 


35. 


Des Lacs National Wildlife 
Refuge, Ward County 


8 May 1968 


A. M. Gammell 


Breeding 


one 


36. 


Fargo, Cass County 


20 December 1969 


Fargo-Moorhead Christmas 


Nonbreeding 


one 








Bird Count (Cruickshank 1970) 




37. 


North of Richardton, Stark County 


July 1970 


J. Hoff 


Breeding 


two -1- young 


38. 


Audubon National Wildlife 
Refuge, McLean County 


8-9 May 1971 


D. C. McGlaughlin 
(Houston 1971) 


Breeding 


one 


39. 


Montpelier, Stutsman County 


11, 13-15 May 1971 


L. C. Haynes (Houston 1971) 


Breeding 


one + one 


40. 


Bowman, Bowman County 


15 May 1971 


L Oberfoel (Houston 1971) 


Breeding 


one 


41. 


Kenmare, Ward County 


3 August 1971 


R. T. Gammell 


Breeding 


one (banded) 


42. 


Gackle, Logan County 


16 May 1972 


G. L. Krapu 


Breeding 


one 


43. 


Jamestown, Stutsman County 


4 June 1972 


M. L. Avery (Stewart 1975) 


Breeding 


one 


44. 


Fargo, Cass County 


12 September 1972; 
30 November 1972 
to 14 April 1973 


A. M. Bolin (Houston and 
Houston 1973), also reported 
on Fargo-Moorhead Christmas 
Bird Count during Count 
week (Arbib et al. 1973) 


Nonbreeding 


one (injured) 


45. 


Bismarck, Burleigh County 


28 May 1973 


R. N. Randall (Houston 1973) 


Breeding 


one 


46. 


Cooperstown, Griggs County 


6 July 1973 


D. L. Kubischta 


Breeding 


one 


47. 


Fargo, Cass County 


8 August 1973 


Observer not specified 

(from the files of the 

North Dakota Birding Society) 


Breeding 


one 


48. 


Bismarck, Burleigh County 


26 May 1974 


B. Quanrud 


Breeding 


one 


49. 


Kulm, LaMoure County 


10 May 1975 


Observer not specified 

(from the files of the 

North Dakota Birding Society) 


Breeding 


one 


50. 


Jamestown, Stutsman County 


18 May 1976 


D. H. Johnson and 

J. E. Johnson 

(Johnson and Johnson 1976) 


Breeding 


one 


51. 


Fargo, Cass County 


25 May 1976 


J. F. Cassel (Serr 1976a) 


Breeding 


one 


52. 


Fargo, Cass County 


22 June to 
11 July 1976 


V.J. Schell(Serr 1976b) 


Breeding 


one 


53. 


Sargent County 


29 June 1977 


R. Schmidt, J. Herman, 
and K. Wilson 


Breeding 


one 


54. 


Hensler, Oliver County 


3 July 1977 


R. N. Randall 


Breeding 


one 


55. 


Fargo, Cass County 


3 November 1977 


C. Brakke 


Nonbreeding 


one 


56. 


Fargo, Cass County 


5-30 November 1977 A. Lies 


Nonbreeding 


one 


57. 


Crosby, Divide County 


19 May 1978 


R. A. Stromstad (Serr 1978) 


Breeding 


one 


58. 


Upper Souris National 
Wildlife Refuge, Ward County 


27 May 1979 


I. 0. Rostad (Serr 1979a) 


Breeding 


one 


59. 


Northern Prairie Wildlife 
Research Center, Stutsman County 


5- 16 July 1979 


C. A. Faanes and other 
Center staff (Serr 1979b) 


Breeding 


one 


60. 


Grand Forks, Grand Forks County 


6-7 May 1980 


J. Van Rybroek (Serr 1980) 


Breeding 


one 


61. 


Oakes, Dickey County 


8 June to 25 


D. Child. J. Fontaine. T. Gat/.. 


Breeding 


two + two nests 






July 1980 


M. Johnson, and J. Oswald 
(Child etal. 1980) 




(totaling 
8 young) 



(Conliniu'd) 



96 



The Canadian Field-Naturalist 



Vol. 116 



Appendix A. (continued) 



No. Location 



Date(s) 



Observer(s) 



Season^ 



Number of birds 



62. Fargo, Cass County 



63. Grand Forks Air Force Base, 
Grand Forks County 



3-18 November 1981 M. A. Bergan and 
C. A. Spurbeck 
(Lambeth 1982) 

19 May 1982 J. F. Kelly (Berkey 1982) 



Nonbreeding one 



Breeding 



64. 


Mercer County 


22 July 1982 


C. A. Faanes (Faanes 1983) 


Breeding 


one 


65. 


Fargo, Cass County 


11 May 1984 


M. A. Bergan and 


Breeding 


one 








C. A. Spurbeck (Berkey 1984) 




66. 


Fargo, Cass County 


21 June 1984 


M. A. Bergan and 
C. A. Spurbeck 


Breeding 


one (injured) 


67. 


Mandan, Morton County 


27 October 1984 


J. Swanick and 0. Swanick 
(Lambeth 1985) 


Nonbreeding 


one 


68. 


Fargo, Cass County 


17 November 1984 


T. Dahlen (Lambeth 1985) 


Nonbreeding 


one 


69. 


Lake Alice, Ramsey County 


2 May 1985 


R. W. Schnaderbeck 


Breeding 


one 


70. 


McLean County 


19 June 1985 


R. E. Martin (Parshall 
Breeding Bird Survey) 


Breeding 


one 


71. 


Hettinger, Adams County 


28-29 April 1986 


D. Griffiths and C. Griffiths 


Breeding 


one 


72. 


Fargo, Cass County 


22 May 1987 


L. L. Falk 


Breeding 


one 


73. 


New Town, Mountrail County 


22-23 May 1987 


R. A. Satermo and 
B. C. Houser 


Breeding 


one 


74. 


Sioux County 


24 May 1987 


R. Titus 


Breeding 


one 


75. 


McHenry County 


21 June 1987 


R. L. Rytter (Denbigh 
Breeding Bird Survey) 


Breeding 


one 


76. 


Grand Forks, Grand Forks County 


Late June 1987 


Observer not specified (from 
the files of the North Dakota 
Birding Society) 


Breeding 


two 


77. 


Eddy County 


5 August 1987 


R. Hill 


Breeding 


one 


78. 


Hettinger, Adams County 


22 April 1988 


D. Griffiths and C. Griffiths 
(Berkey 1988a) 


Breeding 


one 


79. 


J. Clark Salyer National Wildlife 


1 3 May and 


W. Berg, R. E. Martin, 


Breeding 


two 




Refuge, McHenry County 


15 May 1988 


and G. B. Berkey 
(Berkey 1988a) 






80. 


Richardton, Stark County 


June 1988 


J. Hoff 


Breeding 


two + nest 
(with eggs) 


81. 


Fargo, Cass County 


7 June to 
11 July 1988 


R. Kain (Berkey 1988b) 


Breeding 


one 


82. 


Grand Forks, Grand Forks County 


25 October to 

17 November 1988 


D. 0. Lambeth 
(Lambeth 1989) 


Nonbreeding 


one 


83. 


Hettinger, Adams County 


31 May 1990 


D. Griffiths and C. Griffiths 
(Lambeth 1990) 


Breeding 


one 


84. 


Lostwood National Wildlife 
Refuge, Burke County 


9 July 1990 


R. K. Murphy 


Breeding 


one 


85. 


Fargo, Cass County 


10-14 May 1991 


L 0. Hersch (Lambeth 1991) 


Breeding 


one 


86. 


Mercer County 


29 May 1991 


R. E. Martin (Lambeth 1991) 


Breeding 


one 


87. 


Bucyrus, Adams County 


10 June 1991 


D. Griffiths and C. Griffiths 


Breeding 


one 


88. 


Kelly's Slough National Wildlife 
Refuge, Grand Forks County 


23 June 1991 


E. E. Freeberg 


Breeding 


one 


89. 


Fargo, Cass County 


11 May 1992 


L. L. Falk 


Breeding 


one 


90. 


Kidder County 


16 May 1992 


M. A. Otnes 


Breeding 


one 


91. 


Jamestown, Stutsman County 


18 May 1992 


J. T. Lokemoen 


Breeding 


one 


92. 


Bowman County 


24 May 1992 


R. E. Martin and G. B. Berkey 


Breeding 


one 


93. 


Mirror Pool, Ransom County 


19 July 1992 


M. A. Otnes 


Breeding 


one 


94. 


Grand Forks, Grand Forks County 


28 October to 

30 November 1992 


D. 0. Lambeth 


Nonbreeding 


one 


95. 


Fried, Stutsman County 


4 May 1993 


J. T. Price and A. Price 


Breeding 


one 


96. 


Fargo, Cass County 


10-19 May 1993 


D. Wiesenbom 


Breeding 


one 


97. 


South of Bismarck, Burleigh Count) 


15 May 1993 


R. H. O'Connor, 
G. E. Nielsen, and 
M. A. Otnes 


Breeding 


one 












(Continued) 



2002 



Igl and Martin: Mockingbird Occurrences in North Dakota 



97 



Appendix A. (continued) 



No. Location 



Date(s) 



0bser\er(s) 



Season^ 



Number of birds 



98. Sheridan County 



99. Hettinger County 

100. Lostwood National Wildlife 
Refuge. Burke County 

101. Bismarck. Burleigh County 

102. Hettinger. Adams County 

103. Minot Sewage Lagoons, 
Ward County 

104. Bottineau County 

105. Grand Forks, Grand Forks County 

106. Theodore Roosevelt National Park 
(South Unit), Billings County 

107. McHenr\' County 

108. Hettinger. Adams County 

109. Northern Prairie Wildlife 
Research Center. Stutsman County 

1 10. North of Cayuga, Sargent County 

111. Arrowwood National Wildlife 
Refuge. Smtsman County 

1 12. Near Richardton. Stark County 

113. Mandan, Morton County 

1 14. McKenzie County 

115. Arrowwood National 

Wildlife Refuge, Stutsman County 

116. Fargo, Cass County 

1 17. Northern Prairie Wildlife 
Research Center. Stutsman County 

1 1 8. Fargo. Cass County 

1 19. Richardton. Stark County 

120. Mandan, Morton County 

121. Grand Forks County 

122. Hettinger County 

123. Kidder County 

124. New Hradec. Dunn County 

125. Portland. Traill County 

126. Adams County 

127. Southeast of Jamestown, 
Stutsman County 



1 28. Fargo, Cass County 



13 June 1993 

19 June 1993 
27 June 1993 

20 December 1993 
11 May 1994 

18 May 1994 

22 May 1994 
26 May 1994 
5 June 1994 

5-12 June 1994 
13-29 July 1994 
7 September to 



R. E. Martin (Sheyenne Breeding 

Lake Breeding Bird Survey) 
(Berkey 1993) 

D. Griffiths (Bentley Breeding Breeding 
Bird Survey) (Berkey 1993) 

E. M. Madden (Berkey 1993) Breeding 



one 



one 



Bismarck-Mandan Christmas 
Bird Count (LeBaron 1994) 
D. Griffiths and C. Griffiths 
(Martin 1994) 
G. B. Berkey (Martin 1994) 

R. E. Martin (Martin 1994) 
D. O. Lambeth (Martin 1994) 
H. C. Talkington 

R. E. Martin 

D. Griffiths and C. Griffiths 

L. D. Igl D. H. Johnson, 



mid-November 1994 and J. T. Price 



10 May 1995 
13 May 1995 

27 July 1995 



K. Askerooth 
R. Ziamo 

J. Hoff (Berkey 1995) 



15 November 1995 R. N. Randall 
25 November 1995 B. Boyd 
10 May 1996 C.R.Luna 



24 May 1996 
10 June 1996 

28 June 1997 

19 July 1997 

15 April 1998 

16 and 18 May 1998 
1 July 1998 

25 June 1998 

27 October 1998 
30 October 1998 
18 August 1999 
Mid-December 1999 
to 1 February 2000 



9 November 2000 



D. Wiesenbom 
H. A. Kantrud 

C. R. Gjer\'old and 
R. Gjervold 
LHoff 

H. C. Talkington 

J. Maxwell and B. Maxwell 

D. Griffiths (Bentley 
Breeding Bird Survey) 
L. D. Igl (Martin 1998) 

J. Lefor 

C. D. Ellingson 

D. Griffiths and C. Griffiths 
M. A. Sovada and 

R. J. Greenwood; also 
reported on Jamestown 
Christmas Bird Count 
during Count week 
(Randall 2000) 
C. Norheim 



Nonbreedins one 



Breeding 



one 



Breeding 


one 


Breeding 


one 


Breeding 


one 


Breeding 


one 


Breeding 


two 


Breeding 


one 


Nonbreeding 


one 


Breeding 


one 


Breeding 


one 


Breeding 


tw -t- three 




fledglings 


Nonbreeding 


one 


Nonbreeding 


one 


Breeding 


one 


Breeding 


one 


Breeding 


one 



Breeding 

Breeding 

Breeding 
Breeding 
Breeding 

Breeding 

Nonbreeding 
Nonbreeding 
Breeding 
Nonbreedins 



one 

one 
one 
one 
one 

two + nest 

(with four eggs) 

one 

one 

one 

one 



Nonbreeding one 



K)bservations between 15 April and 31 August were considered breeding-season records, and obscr\aiions bctueen 1 September and 
14 April were considered nonbreeding-season records. 



Diets of Northern Flying Squirrels, Glaucomys sabrinus, 
in Southeast Alaska 

Sanjay Pyare', Winston P. Smith^ Jeffrey V. Nicholls^, and Joseph A. Cook^"^ 

'Denver Zoological Foundation, P.O. Box 20377, Juneau, Alaska 99802 USA [corresponding author] 

-United States Forest Service, Pacific Northwest Region, Forestry Sciences Laboratory, 2770 Sherwood Lane, Suite 2A, 

Juneau, Alaska 99801 USA 
-^University of Alaska Museum, Fairbanks, Alaska 99775-6960 USA 
■^Current address: Department of Biological Sciences, Idaho State University, Pocatello, Idaho 83209-8007 USA 

Sanjay Pyare, Winston P. Smith, Jeffrey V. Nicholls, and Joseph A. Cook. 2002. Diets of Northern Flying Squirrels, 
Glaucomys sabrinus, in southeast Alaska. Canadian Field-NaturaUst 116(1): 98-103. 

We examined the diet of the Northern Flying Squirrel {Glaucomys sabrinus) during summer and autumn seasons in tem- 
perate rain-forest habitat of Southeast Alaska, a region in which the ecology of this species is poorly understood. Truffles, a 
food item that is commonly consumed by squirrels during snow-free periods outside of Alaska, were present less frequent- 
ly in squirrel feces than two other food items, epigeous fungi and vegetation, although no food item dominated fecal com- 
position. Truffles were less frequent in fecal samples from mixed-conifer muskeg habitats than from old-growth forest 
habitats. Overall, we found that squirrels consumed a total of five truffle genera; Elaphomyces and Hymenogaster being the 
most common. Compared to populations in the western contiguous United States, squirrel populations in Southeast Alaska 
consumed truffles less frequently and consumed a smaller total number of truffle genera. In addition, samples from individ- 
ual squirrels in Alaska tended to contain fewer genera than samples from the contiguous United States. Finally, squirrels in 
Alaska consumed other food items such as vascular vegetation, lichens, and mushrooms more frequently than squirrels in 
other geographic areas. These patterns suggest that the association between flying squirrels and truffles may be relatively 
weaker in Southeast Alaska than has been documented elsewhere. Consequently, additional information on life history and 
ecology of flying squirrels is warranted before forest management guidelines can be developed. 

Key Words: Alaska, diet, lichens, mycophagy. Northern Flying Squirrels, old-growth. Pacific Northwest, truffles 



Northern Flying Squirrels (Glaucomys sabrinus) 
are mycophagous and frequently feed on the fruiting 
bodies of hypogeous fungi (i.e., truffles) in conifer- 
ous forests of western North America (Carey et al. 
2000; Colgan et al. 1997; Hall 1991; Maser et al. 
1985; Maser et al. 1986; Maser and Maser 1988; 
Pyare and Longland 2001; Rosentreter et al. 1997; 
Waters and Zabel 1995). Because spores of these 
fungi are mycorrhizal and remain viable after pas- 
sage through small mammal digestive tracts, flying 
squirrels are thought to play an important ecological 
role by dispersing these spores (Cazares and Trappe 
1990; Cork and Kenagy 1989; Kotter and Farentinos 
1984; Li et al. 1986; Maser and Maser 1988). 

During winter, truffles are uncommon food items 
for flying squirrels because truffles are less abundant 
and/or snow cover may limit access to these below- 
ground fungi. Thus, alternative and readily available 
food items, such as lichens, are predominant winter 
food items for flying squirrels. During snow-free 
periods, however, such as in summer and autumn, 
flying squirrels are thought to specialize on truffles 
because flying-squirrel stomachs and feces generally 
contain a diverse array of truffle genera. Furthermore, 
squirrels exhibit preferences for truffles over other 
food items (Zabel and Waters 1997). For the most 
part, however, our knowledge about these squirrels is 
restricted to geographic areas where these fungi are 



abundant and where fungal composition is relatively 
well documented, particularly in southern regions of 
the Pacific Northwest, such as in northern California, 
Oregon, and Washington, U.S.A. (Carey et al. 2000; 
Colgan et al. 1997; Hall 1991; Maser et al. 1985; 
Maser et al. 1986; Maser and Maser 1988; Pyare and 
Longland 2001; Waters and Zabel 1995). In contrast, 
in northern regions of the Pacific Northwest, such as 
Southeast Alaska and adjacent British Columbia, lit- 
tle is known about the ecology and food habits of fly- 
ing squirrels, nor is much known about the distribu- 
tion of truffles. This is true despite that coastal and 
interior forests of this region provide extensive habi- 
tat for flying squirrels, and provide an abundance of 
conifers that potentially serve as ectomycorrhizal 
hosts for truffle fungi. In general, however, it is 
thought that truffles are relatively uncommon in 
northern forests possibly because climatic conditions 
may be too extreme and/or this region may simply 
represent the northern limits of truffle distribution for 
some species (E. Cazares, personal communication). 
Thus, even during snow-free periods, truffles may be 
less common in this region, and consequently, flying 
squirrels may be more likely to exhibit generalist- 
type feeding behavior there (Thysell et a. 1997). 

Basic information about squirrel ecology and food 
habits in this region would have practical merit for 
contemporary objectives regarding the conservation 



98 



4 



2002 



Pyare, Smith, Nicholls, and Cook: Diets of Flying Squirrels in Alaska 



99 



Alexander Archipelago 




Prince of WaleSr 
Island Complex\ 





ivr^ 






^7^' 



^ 



G\ 






1-:^ 











Figure 1. Location of study area (Prince of Wales Island 
complex) in the southern Alexander Archipelago of 
Southeast Alaska, U.S.A. Numbered locations are 
as follows: (1) Prince of Wales Island, (2) Heceta 
Island, (3) El Capitain Island, (4) Tuxekan Island, 
and (5) North Island. 



of the flying squirrel. In addition, an endemic sub- 
species, the Prince of Wales Northern Flying 
Squirrel (G. s. griseifrons), is found in Southeast 
Alaska (Demboski et al. 1998; McDonald and Cook 
1996). Thus, to provide a more comprehensive 
understanding of flying-squirrel ecology, we evaluat- 
ed diet patterns of flying squirrels in temperate rain- 
forest habitats of Southeast Alaska during snow-free 
periods and compared these patterns to those in the 
western contiguous United States. 

Methods and Materials 

We conducted microhistological analyses of tly- 
ing-squirrel feces collected from a total of 13 forest 
stands in the Prince of Wales Island complex in the 
southern Alexander Archipelago of Alaska (Figure 
1). These coastal study sites ranged from 2()()-8()() m 



in elevation. Ten stands consisted of old-growth (un- 
logged) forest habitat, six of which were on Prince of 
Wales Island (stands separated by > 5 km): Dargun 
Point (55°54'44" N, 133°15'36" W), Honker 1 
(55°20'48" N, 132°50'08" W), Honker 2 (55°21'02" 
N, 132''50'36), Honker 3 (55°19'54" N, 132°49'37" 
W), Naukati Bay (55°51'25" N, 133°10'02" W) and 
Tuxekan Village (55°54'06" N, 133°14'55" W). An 
additional four old-growth stands were on El Capitan 
Island (55°55'59" N, 133°18'33" W), Heceta Island 
(55M6'00"N, 133°27'00"W), North Island 
(55°57'00"N, 133°19'00"W), and Tuxekan Island 
(55°51'11"N, 133°13'39"W). Finally, three stands 
consisting of mixed-conifer muskeg habitat were on 
Prince of Wales Island (stands separated > 5 km): 
Ball's Lake (55M3'00"N, 132°49'00"W), Control 
Lake 1 (55M3'48"N, 132°48'54"W), and Control 
Lake 2 (55°43'24", 13249'39" W). 

Old-growth habitat included Sitka Spruce (Picea 
sitchensis). Western Hemlock {Tsuga heterophylla) 
and Red {Thuja plicata) and Yellow cedar 
(Chamaecyparis nootkatensis). Large trees, downed 
and decaying wood, snags and small gaps created by 
individual blow-downs all were key components of 
this habitat. Dense patches of Blueberry (Vaccinium 
spp.) dominated the understory. In contrast, mixed- 
conifer muskeg habitat exhibited poor drainage, shal- 
low organic soils, and only patches of mixed conifer 
vegetation that occurred on gently sloping lowlands 
and floodplains. Conifer vegetation included Yellow 
Cedar, Red Cedar, Western Hemlock, Mountain 
Hemlock (T. mertensiana). Shore Pine {Piniis contor- 
ta var. contorta), and Sitka Spruce. Understory vege- 
tation varied considerably within this habitat type. 
Open areas with little overstory consisted of a mix- 
ture of herbaceous species (sedges, grasses. Skunk 
Cabbage [Lysichitum americanum]) and Labrador 
Tea {Ledum glandulosum), whereas Blueberry domi- 
nated areas with well-developed overstories. Because 
muskegs are a previously undocumented habitat for 
Northern Flying Squirrels, we treated data from these 
stands separately. 

We collected fecal samples from flying squirrels in 
these habitats only during snow-free periods; i.e., 
when truffles were most likely to be available and/or 
snow could not preclude access to truffles. On three 
of the old-growth sites (Honker 1-3) and all three of 
the muskeg sites, fecal samples were collected from 
the anus of individually-marked squirrels that were 
live-trapped in Tomahawk live-traps during autumn 
(September-October) of 1999 and 2000. No more 
than one sample was collected from each individual. 
We did not collect samples that appeared to consist of 
bait (a peanut butter-rolled oats mixture). For the 
remaining seven old-growth sites, feces were collect- 
ed from the digestive tract o{ museum specimens 
(University of Alaska Museum, Fairbanks, Alaska). 
These samples were originally collected during sum- 
mer (June-July) and autumn 1999. 



100 



The Canadian Field-Naturalist 



Vol. 116 



Table 1. Prevalence of different food items in fecal samples collected from flying squirrels in old-growth stands (n = 10) 

and muskeg stands (3) in Southeast Alaska. Truffle genera found in each stand are shown at right. 









% Occurrence 










N 


Truffles 


Lichen 


Vegetation 


Mushrooms 


Insect 


Truffle Genera Present 


Old Growth Stand 
















Honker 1 


34 


50 


50 


50 


59 


12 


Elaphomyces, Hymenogaster 


Honker 2 


14 


50 


20 


23 


30 





Sclerogaster 


Honker 3 


21 


100 


20 


56 


20 





Elaphomyces 


Dargan Point 


6 


50 


12 


50 








Hymenogaster 


Naukati 


5 


64 


20 


75 


48 


4 


Elaphomyces, Hymenogaster 


Tuxekan Village 


10 


20 


64 


24 


40 





Elaphomyces 


Heceta Island 


17 


40 


10 


64 


23 


12 


Elaphomyces, Hymenogaster 


Tuxekan Island 


26 


36 


36 


60 


24 


12 


Elaphomyces 


North Island 


3 


12 


30 


84 


33 


4 


Elaphomyces 


EI Capitan Island 


14 


82 


8 


56 


84 





Hymenogaster 




X 


50.4 


27 


55.2 


36.1 


4.4 






s.d. 


26.8 


18.4 


20.1 


23.3 


4.6 




Muskeg Stands 
















Ball's Lake 1 


4 


4 


20 


60 


50 





Elaphomyces 


Control Lake I 


5 


12 


40 


28 


40 





Elaphomyces 


Control Lake 2 


8 


40 





50 


88 





Elaphomyces 




X 


18.7 


20 


46 


59.3 


0.0 






s.d. 


15.5 


16.3 


12.5 


21.6 


0.0 





All samples were stored separately in glass vials 
with 70% ethanol. For analysis, small portions of 
each sample were mixed with 3ml of distilled HjO, 
and mixed vigorously together to achieve a homoge- 
neous solution. A small drop of the solution was 
mixed with a drop of Melzer's reagent and placed 
on a slide, covered with a 18 mm circular slide 
cover, and observed using bright-field microscopy 
at 40 X . We identified the following different food 
items in 25 randomly selected field-of-views: the 
spores of hypogeous fungi and epigeous fungi, algal 
cells from lichen, and remains of vascular vegeta- 
tion and insects. Spores of hypogeous fungi were 



further identified to genus using a spore key 
(Castellano et al. 1989). 

Results 

We analyzed a total of 151 fecal samples from 
old-growth forest sites, and 17 samples from muskeg 
sites. Among old-growth samples, vascular vegeta- 
tion was the most frequent food item (x± se = 
55 ± 6.4%), while truffle spores were second most 
frequent (50 ± 8.5%) (Table 1). In contrast, among 
muskeg samples, both epigeous fungi and vegetation 
items were more frequent (59 ± 12.5% and 
46 ± 7.2%, respectively) than truffles (19 ± 8.9%). 



Table 2. Truffle-consumption patterns of various flying-squirrel populations in western North America. All data were 
collected in mature conifer-forest habitats. Seasons correspond to the following months: Spring (March-May). Summer 
(June-August). Autumn (September-November), and Winter (December-February). 



Location and Source 



N 



9c Occurrence Bv Season 



Spring 



NevadaCounty. California: Hall 1991 107 92 

Lake Tahoe Basin, California: Pyare and Longland 2001 93 

Lassen County. California: McKeever 1960 24 83 

Plumas County, California: Waters and Zabel 1995 88 

Douglas County. Oregon: Maseret al. 1986 162 97 

Douglas and Coos County, Oregon: Carey et al. 1999 158 75 

Lane and Linn County, Oregon: Maser et al. 1985 28 

Blue Mountain Province, Oregon: Maseret al. 1985 63 

Watson Falls, Oregon: Ca/.ares et al. 1999 12 

Payette National Forest, Idaho: Rosentreter et al. 1997 200 77 

Thurston County, Washington: Colgan et al. 1997 12 

Southeast Alaska (this study) 151 



Summer Autumn 



92 
100 
100 
100 

93 



94 

78 

50 



81 
100 
100 

99 

100 
100 
100 



50 



Winter 



81 
56 
98 



77 
85 



2002 



Pyare, Smith, Nicholls, and Cook: Dets of Flying Squirrels in Alaska 



101 



100- 



0) 

o 

c 

2 

3 
O 
U 

O 




■ Alaska 


n Non- 


Alaska 



/^ ^^ ^^^ o^^"^ 
Food Type 

Figure 2. Comparison of food-item occurrence in pooled 
samples between southeast Alaska (n = 10 stands) 
and outside Alaska (9). Insect frequencies were not 
reported in most studies and therefore left out of 
this comparison. All samples were collected in 
mature conifer-forest habitat during snow-free peri- 
ods. Bars represent standard error. Non- Alaska data 
from: Cazares et al. 1999; Hall 1991; Maser et al. 
1985; Maser et al. 1986; McKeever 1960; Pyare and 
Longland 2001; Rosentreter et al. 1997; and Waters 
and Zabel 1995. 



The frequency of lichen in samples was roughly simi- 
lar between muskeg and old-growth sites (20 ± 9.4% 
and 27 ± 5.8%, respectively). Five truffle genera were 
present in old-growth samples. Elaphomyces was the 
only truffle genus present in muskeg samples. 

Examination of spring and autumn fecal-analysis 
data from 1 1 other forest sites throughout the western 
United States revealed that overall, the frequency of 
truffles in fecal samples from southeast Alaska was 
comparatively low (Table 2; Figure 2). In contrast, 
the frequencies of lichens, vascular plants, and epi- 
geous fungi were higher in Southeast Alaska than in 
the contiguous western United States, although these 
differences were not significant (Figure 2). 

Although sample size had a strong influence on the 
number of truffle genera present in samples collected 
throughout the western contiguous United States 
(Figure 3), Southeast Alaska appears to be atypical 
and does not fit this general correlation. Taxonomic 
richness of Alaska diets was much lower than other- 
wise would have been predicted (roughly 22 genera) 
for the observed Alaska sample size (n = 151; old- 
growth habitat only). On average, individual squirrel 
samples from Alaska contained fewer genera than 
samples from California and Oregon (Figure 4). 

Discussion 

Although flying squirrels are mycophagous 
throughout their range, dietary patterns of popula- 



tions in Southeast Alaska appeared to differ from 
those in the western contiguous United States during 
snow-free periods. Compared to their southern coun- 
terparts, at the population level, squirrels in 
Southeast Alaska consumed truffles less frequently 
and consumed a smaller total number of truffle gen- 
era. Furthermore, individual squirrel samples from 
Alaska contained fewer genera than southern fecal 
samples. These patterns suggest that flying squirrels 
may be less dependent on truffles in Southeast 
Alaska, even during seasons when potential avail- 
ability is greatest. This is consistent with other stud- 
ies that have shown that the species may in some 
instances exhibit generalist-type feeding behavior. 
For example. Thy sell et al. (1997) observed wild 
squirrels feeding on other types of highly digestible 
food items such as seeds and berries, and conse- 
quently concluded that previous studies involving 
fecal-analyses have typically underestimated the 
importance of such food items. More recently, 
Claridge et al. (1999) concluded that hypogeous 
fungi were of moderate nutritional value to captive 
flying squirrels, which failed to gain body mass 
when fed solely the common fungal sporocarp 
Rhizopogon vinicolor. Additional support for the 
idea that flying squirrels may in some part of their 
range be only marginally dependent on truffles 
comes from the southern Appalachian Mountains, 
where truffle abundance is relatively low and only a 
few common truffle genera are found in fecal sam- 
ples (Loeb et al. 2000; Weigl et al. 1992). 



<30 - 

so- 










♦ 


ls 

0) 25- 

c 

0) 

e> 20- 








♦ 






ii5. 

H 

d 10- 

z 


♦ 
♦....••••••■■■■■■■" 


...••••■■*"' 


•■■■■■'♦ 

♦ 


SEA 


lash 


5- 








♦ 




0- 








1 


— 1 



50 



100 



150 



200 



No. Fecal Samples Analyzed 



Figure 3. Relationship between sample size and total num- 
ber of triiftle genera detected in fecal samples of 
flying-squirrel populations in western North 
America (r = 0.90 not including Alaska data). Non- 
Alaska data from: Carey et al. 1999; Ca/ares et al. 
1999: Colgan et al. 1997; Hall 1991; Maser et al. 
1983; Maser et al. 1986; Pyare and Longland 2(X)I; 
and Rosentreter et al. 1997. 



102 



The Canadian Field-Naturalist. 



Vol. 116 




Location 

Figure 4. Comparison of the mean number of truffle genera 
per diet sample (i.e. per individual) among four fly- 
ing-squirrel populations. All data were collected 
during snow-free periods in mature conifer-forest 
habitat. Sample sizes and data sources are as fol- 
lows: Alaska (this study, n = 151); California (Pyare 
and Longland 2001, 93); NE Oregon (Maser et al. 
1985, 63); and NW Oregon (Maser et al. 1985, 28). 



Although we did not assay truffle abundance at 
these study sites, and virtually nothing is known 
about truffle distribution in Southeast Alaska, the 
simplest explanation for the disparity in squirrel 
diets between Southeast Alaska and elsewhere may 
be due to low truffle availability. Alternatively, 
snow cover could not have been a factor for the rel- 
atively low consumption rates of truffles observed 
in our study. Nor is it likely that the disparity is 
explained by differences in food preferences, 
although we cannot exclude this as a possibility. 
We never encountered truffles at these study sites, 
nor did we ever witness mycophagous animal dig- 
gings, clues that both scientists and hobbyists com- 
monly use to locate these subterranean fruiting bod- 
ies (Waters and Zabel 1995). In contrast, S. P. fre- 
quently observed evidence of the association 
between these animal diggings and truffles in 
mixed-conifer forest of the Sierra Nevada Range, 
California. In samples from both old-growth and 
muskeg habitats in Alaska, the preponderance of 
Elaphomyces spp. further suggests that fungal 
diversity may be limited perhaps due to adverse cli- 
matic conditions of this cool, extremely wet region. 
Elaphomyces spp. have a relatively thick ( 0.5 cm) 
outer peridium and are thought of as being among 
the hardiest of hypogeous fruiting bodies. We are 
particularly skeptical that truffles fruit to any sig- 



nificant degree in the more open areas of muskeg 
habitats, the soil of which is often poorly drained 
and flooded. 

The frequency of lichen, vegetation, and mush- 
rooms were all higher in Southeast Alaska samples 
than samples from elsewhere, although differences 
were not significant. These other food items may be 
relatively more important than our simple fecal pel- 
let analysis revealed; volume estimates of stomach 
contents would certainly have provided a better 
understanding of ingestion rates of different food 
items. Of particular relevance to flying squirrels in 
this region may be the abundance of arboreal 
lichens, which achieve high biomass and diversity. 
In particular, Bryoria spp. and Usnea spp. are known 
to be consumed by flying squirrels (Rosentreter et al. 
1997; Zabel and Waters 1997) and appear to be pre- 
ferred over other lichens by captive animals (S. 
Pyare, unpublished data). We commonly observed 
both of these lichens in these old-growth forest and 
muskeg habitats during our three-year study of fly- 
ing squirrels in Alaska. 

Conservation and management measures for fly- 
ing squirrels have conventionally emphasized the 
underlying importance of hypogeous fungi (Loeb et 
al. 2000; Maser and Maser 1988; Zabel and Waters 
1997). Management plans in Southeast Alaska and in 
other northern regions, such as coastal and interior 
British Columbia, however, may further need to rec- 
ognize the complete array of food items consumed 
by these northern populations. For instance, in the 
western contiguous United States, among studies of 
the impact of disturbances, such as fire, clear cutting, 
and selective harvesting, on flying squirrel popula- 
tions, there have typically been concomitant efforts 
to assay the impacts on truffle abundance or diversi- 
ty, while the impacts of disturbance on other food 
types have not been evaluated (Amaranthus et al. 
1994; Carey et al. 2000; Cazares et al. 1999; Colgan 
1997; Waters et al. 1994; Waters and Zabel 1995). 
Management strategies in northern regions, however, 
may require a more pluralistic understanding of dis- 
turbance impacts on lichens, vascular plants, and 
mushrooms, as well as belowground fungi. 
Furthermore, because patterns of foraging, move- 
ment, and habitat use differ as a consequence of 
these different feeding habits, management strategies 
may require additional efforts to understand the 
unique ecological niche flying squirrels occupy in 
this particular region. 

Acknowledgments 

Aren Eddingsaas, John Frisch, Jeff Hayes, Meade 
Krosby, Kari Murabito, and Kevin White diligently 
helped obtain diet samples of flying squirrels in 
Southeast Alaska. Collin Gillin of Tufts University 
kindly furnished laboratory space and a microscope 
for diet analyses. Efren Cazares and Jim Trappe had 



2002 



Pyare, Smith, Nicholls, and Cook: Diets of Flying Squirrels in Alaska 



103 



previously provided their expertise with fungal iden- 
tification at Oregon State University, Corvallis. This 
study was supported by the University of Alaska 
Museum, Fairbanks, Alaska, and the USDA Forest 
Service's Forestry Sciences Lab in Juneau, Alaska. 
S. Pyare was supported by the Wildlife Conservation 
Society and the Denver Zoological Foundation dur- 
ing drafting of this manuscript. 

Literature Cited 

Amaranthus, M., J. M. Trappe, L. Bednar, and D. 
Arthur. 1997. Hypogeous fungal production in mature 
Douglas-fir forest fragments and surrounding plantations 
and its relation to coarse woody debris and animal 
mycophagy. Canadian Journal of Forest Research 24: 
2157-2165. 

Carey, A. B., J. Kershner, B. Biswell, and L. A. 
Dominguez de Toledo. 2000. Ecological scale and for- 
est development: squirrels, dietary fungi, and vascular 
plants in managed and unmanaged forests. Wildlife 
Monographs 142: 1-71. 

Castellano, M., J. M. Trappe, Z. Maser, and C. Maser. 
1989. Keys to the spores of hypogeous fungi of northern 
temperate forests with special reference to animal 
mycophagy. Mad River Press, Eureka, California. 186 
pages. 

Cazares, E., D. L. Luoma, M. P. Amaranthus, C. L. 
Chambers, and J. F. Lehmkuhl. 1999. Interaction of 
fungal sporocarp production with small mammal abun- 
dance and diet in Douglas-fir stands of the southern 
Cascade Range. Northwest Science 73: 64-76. 

Cazares, E., and J. M. Trappe. 1990. Spore dispersal of 
ectomycorrhizal fungi on a glacier forefront by mammal 
mycophagy. Oecologia 86: 507-510. 

Claridge, A., J. M. Trappe, S. J. Cork, and D. L. Claridge. 
1999. Mycophagy by small mammals in the coniferous 
forests of North America: nutritional value of sporocarps 
of Rhizopogon vinicolor, a common hypogeous fungus. 
Journal of Comparative Physiology B 169: 172-178. 

Colgan, W. 1997. Diversity, productivity, and mycophagy 
of hypogeous mycorrhizal fungi in a variably thinned 
Douglas-fir forest. Ph.D. dissertation. Oregon State 
University, Corvallis, Oregon. 

Colgan, W., A. B. Carey, and J. M. Trappe. 1997. A reli- 
able method of analyzing dietaries of mycophagous 
small mammals. Northwestern Naturalist 78: 65-69. 

Cork, S. J., and G. J. Kenagy, 1989. Rates of gut passage 
and retention of hypogeous fungal spores in two forest 
dwelling rodents. Journal of Mammalogy 70: 512-519. 

Demboski, J. R., B. K. Jacobsen, and J. A. Cook. 1998. 
Endemism in the Alexander Archipelago: an assessment 
of genetic variation in northern flying squirrels 
(Rodentia: Glaucomys sabrinus). Canadian Journal of 
Zoology 76: 1771-1777. 

Hall, D. S. 1991. Diet of the northern flying squirrel at 
Sagehen Creek, California. Journal of Mammalogy 72: 
615-617. 



Kotter, M. M., and R. C. Farentinos. 1984. Formulation 
of ponderosa pine ectomycorrhizae after inoculation 
with feces of tassel-eared squirrels. Mycologia 76: 
758-760. 

Li, C. Y., C. Maser, Z. Maser, and B. Caldwell. 1986. 
Role of three rodents in forest nitrogen fixation in west- 
ern Oregon: another aspect of mammalian-fungus-tree 
mutualism. Great Basin Naturalist 46: 41 1^14. 

Loeb, S. C, F. H. Tainter, and E. Cazares. 2000. Habitat 
associations of hypogeous fungi in the southern Appala- 
chians: Implications for the endangered northern flying 
squirrel {Glaucomys sabrinus coloratus). American 
Midland Naturalist 144: 286-296. 

Maser, C, and Z. Maser. 1988. Interactions among squir- 
rels, mycorrhizal fungi, and coniferous forests in Oregon. 
Great Basin Naturalist 48: 358-369. 

Maser, C, Z. Maser, J. W. Witt, and G. Hunt. 1986. The 
northern flying squirrel: a mycophagist in Oregon. 
Canadian Journal of Zoology 64: 2086-2089. 

Maser, Z., C. Maser, and J. M. Trappe. 1985. Food habits 
of the northern flying squirrel in Oregon. Canadian 
Journal of Zoology 63: 1084-1088. 

McDonald, S. O., and J. A. Cook. 1996. The land mammal 
fauna of southeast Alaska. Canadian Field-Naturalist 
110:571-598. 

McKeever, S. 1960. Food of the northern flying squirrel in 
northeastern California. Journal of Mammalogy 41: 
270-271. 

Pyare, S., and W. S. Longland. 2001. Patterns of ectomyc- 
orrhizal fungi consumption in remnant old-growth 
forests of the Sierra Nevada. Journal of Mammalogy 82: 
681-689. 

Rosentreter, R., G. D. Hayward, and M. Wicklow- 
Howard. 1997. Northern flying squirrel seasonal food 
habits in the interior conifer forests of central Idaho, 
U.S.A. Northwest Science 71: 97-101. 

Thysell, D. R., L. J. Villa, and A. B. Carey. 1997 
Observations of northern flying squirrel feeding behav- 
ior: use of non-truffle food items. Northwestern 
Naturalist 78: 87-92. 

Waters, J. W., K. S. McKelvey, C. J. Zabel, and W. W. 
Oliver. 1994. The effect of thinning and broadcast burn- 
ing on sporocarp production of hypogeous fungi. 
Canadian Journal of Forest Research 24: 15 16-1522. 

Waters, J. W., and C. J. Zabel. 1995. Northern flying 
squirrel densities in fir forests of northeastern California. 
Journal of Wildlife Management 59: 858-866. 

Weigl, P. D., T. W. Knowles, and A. C. Boynton. 1992. 
The distribution and ecology of the northern flying 
squirrel in the Southern Appalachians. U.S. Fish and 
Wildlife Service. 140 pages. 

Zabel, C. J., and J. R. Waters. 1997. Food preferences of 
captive northern flying squirrels from the Lassen 
National Forest in northeastern California. Northwest 
Science 71: 103-107. 

Received 18 April 2001 
Accepted 18 March 2002 



104 



The Canadian Field-Naturalist 



Vol. 116 



Nesting Activities of an Eastern Spiny Softshell Turtle, 
Apalone spinifera 

Claude Daigle^, Patrick Galois2, and Yves Chagnon^ 

'Societe de la Faune et des Pares du Quebec, direetion de la recherche sur la faune, 675 Rene-Levesque est, Quebec, 

Quebec GIR 5V7 Canada ; e-mail: claude.daigle@fapaq.gouv.qc.ca 
2Societe d'Histoire Naturelle de la Vallee du Saint-Laurent, 21 125 chemin Sainte-Marie, Sainte-Anne-de-Bellevue, 

Quebec H9X 3Y7 Canada; e-mail: pagalois@aei.ca 
^Societe de la Faune et des Pares du Quebec, direction regionale de la Monteregie, 201 place Charles-Lemoyne, Longueuil, 

Quebec J4K 2T5 Canada 

Daigle, Claude, Patrick Galois, and Yves Chagnon. 2002. Nesting activities of an Eastern Spiny Softshell Turtle, Apalone 
spinifera. Canadian Field-Naturahst 1 16(1): 104-107. 

Une tortue-molle a epines a ete suivi etroitement durant la periode de ponte de 1998. Cette tortue a demontre un patron 
quotidien d'activite regulier avant la ponte. Elle consacrait ses matinees a s'exposer au soleil et a se deplacer dans un 
secteur restreint, ses apres-midi a se deplacer sur des distances de 2 a 3 km, pour finalement s'arreter en fin d'apres-midi 
alors que durant la soiree, elle semblait etre a la recherche d'un endroit propice pour deposer ses oeufs. Cette tortue-molle a 
epines a demontre la capacite de se deplacer a contre courant sur une distance de plus de 7 km, en trois jours, pour aller 
pondre. 

An Eastern Spiny Softshell Turtle was closely followed during the 1998 nesting season. It showed a rather consistent daily 
pattern of activity. The morning was spent basking and moving around in a small area, large movements (2-3 km) were 
made during the afternoon and the turtle would stop for the evening, acting like it was searching for a suitable nesting site. 
This softshell turtle moved 7 km upstream in three days for nesting. 

Key Words : Eastern Spiny Softshell Turtle, Apalone spinifera, nesting, movement, activity, Quebec, Canada. 



The information reported here was collected in 
the course of a larger study designed to help the con- 
servation of a unique Quebec population of Eastern 
Spiny Softshell Turtle {Apalone spinifera) (Galois, 
1999*). Attempts were made to follow three females 
to their nesting sites, but we were successful with 
only one. This adult female (535) was captured 18 
June 1997 in Riviere aux Brochets (Figure 1), a trib- 
utary of Lake Champlain (73°13'W, 45°04'N) and 
equipped with a radio transmitter (frequency 
155.535; Holohil System Ltd, Ottawa, Canada) pro- 
vided with a mortality option that produces a double 
bip when the turtle remains still for more than four 
hours. The softshell spent the winter of 1998 in the 
Missisquoi River near Swanton, Vermont, U.S.A. 
(73°13'W, 44°58'N), moved north across Lake 
Champlain during the first half of May, and up in the 
Riviere aux Brochets, where it spent the following 
three weeks in the vicinity of a small island, more 
than 25 km away from its wintering area. We fol- 
lowed 535 for most of the daylight hours during the 
few days before nesting and the day after nesting in 
June 1998. Although nesting activities had been 
described before (Eigenman 1896; Newman 1906; 
Evermann and Clark 1920; Minton 1972; Ernst et al. 
1994), few individuals were involved in those obser- 
vations, and daily activities shortly before and after 
ovoposition has not been reported previously. 



9 June, mostly sunny, 26°C 

At our arrival (10:15), 535 was basking on the 
sunny side of the river, approximately 20 cm from 
water, with two other softshell females basking a 
few meters away. They started moving at around 
11:30, going in and out of the water. By 13:30, 535 
had left the area and moved upstream throughout the 
afternoon. At 18:00, after swimming about 2 km, it 
stopped and remained in the area until we left at 
20:00. 

10 June, sunny, 22° C 

When we arrived at 5:30, 535 was in the same 
area as the previous evening. The signal became 
double at 9:30, indicating that the turtle had not 
moved for the last 4 hours. At 9:30, we spotted 535 
basking on a stone in the river, approximately 5 m 
from the shore. It entered the water at 10:30 but 
remained in the area. By noon, it had started moving 
upstream. The river was about 30 cm deep and 30 m 
wide, and steep banks allowed us to often have a 
good look in the water. On many occasions we could 
see 535 passing by. It swam vigorously, its neck 
fully extended, pointing its nose out of the water 
every 5 to 10 m and sometimes stopping in this posi- 
tion for a few seconds. The softshell kept this pace 
for the whole afternoon. At 17:00, it stopped next to 
a gravel bar 5 m long but only 10 to 20 cm above the 



104 



2002 



Daigle, Galois, and Chagnon: Nesting of Softshell 



105 



ya^oe'oo 



73°04'00 



73''02'00 




La Grande 6a/e 
(LicChmplain} 



73°06'00 



73°04'00 



73°02'00 



FiCiURH 1. Movemcnls ola female Hastern Spiny Sollshell Turtle during the nesting season. 



106 



The Canadian Field-Naturalisx 



Vol. 116 



water level, almost 3 km from its point of departure 
at noon. The softshell came out of the water at 17:30, 
moving around on the gravel bar and rubbing its chin 
on the substrate every now and then. It re-entered the 
water 10 minutes later, but climbed back on it at 
19:50, departing about 20 seconds later when dis- 
turbed by a man passing on the road 10 m away. At 
20:10, it climbed once more on the gravel bar, but 
was disturbed a second time by human activities. 
The softshell was still in the area when we left at 
21:00. 

11 June, mostly sunny, 22° C 

We returned to the area at 6:00. Although not visi- 
ble, 535 was still around. At 8:30, the signal became 
double and remained so until 10:15. For the next 
hour, the turtle seemed to be moving around, but 
staying in the area. At 1 1:15, 535 walked on the grav- 
el bar in spite of the presence of six ducks. Again, the 
softshell seemed to be rubbing its chin on the ground. 
By 11:30, it was back in the water and swam inten- 
sively upstream throughout the afternoon. It climbed 
on the sunny shoreline of the river at 15:50, approxi- 
mately 2.5 km away from its daily departure point. It 
moved in and out of the water on a 30 m stretch of 
the river, next to a sharp curve. At 16:05, 535 walked 
on land for the sixth time, at the other end of the 
curve, about a hundred meters upstream from where 
it had first gone on land, but only 20-30 m in straight 
line through forest. The softshell started the excava- 
tion at 16: 15 at the edge of the vegetation, into gravel 
and flat pebbles ranging in size from 1 to 10 cm long. 
The first part of the digging motion was slow, but it 
ended vigorously and pebbles were thrown up to 2 m 
away. The excavation stopped at 16:35. After a few 
minutes without moving (probably laying), the turtle 
started filling the hole at 16:45. The softshell ran 
back into the water at 16:49, 44 minutes after coming 
out of the water. The nest was excavated 2 m from 
the water and approximately 1 m above the water 
level, in full late afternoon sunshine (northwestern 
slope). Numerous other signs of nesting activity were 
present in the vicinity. The softshell immediately 
started swimming upstream. We observed 535 at 
20:00, 1 km upstream from where it nested, about 
100 m below a dam in a small village. We left the 
area for the night. 

12 June, partially sunny, 24° C 

When we arrived at 6:40, the turtle had moved 
downstream about 0.5 km from where we last saw it 
the night before. The signal was double but became 
single 10 minutes later, indicating that the turtle just 
started to move after a stop of at least four hours. 
The radio indicated that the softshell was moving 
downstream. It swam by its nest at 7:00. A large 
Snapping Turtle {Chelydra serpentina) was digging 
a few meter from where 535 had excavated its nest 
the night before. We saw the softshell at 8:40, and 



then again at 10:15. It was swimming vigorously, in 
the same way it had been during the few days it 
spent moving up the river. We stopped tracking the 
turtle around 11:00 to check the softshell 's nest to 
confirmed that it had really laid eggs. We found eggs 
exactly where the turtle dug the night before. In 
order to avoid any disturbance in the incubation pro- 
cess, eggs were covered, without counting them, as 
soon as we discovered the first egg. We tracked the 
turtle during the whole afternoon and by 17:30, 535 
was back in the area it left 9 June. Travelling down- 
stream, it took the turtle one day to cover the same 
distance that had required three days moving 
upstream. Swimming time summed up to 11 hours 
travelling downstream and 17 hours moving 
upstream. 

Discussion 

Most events observed while tracking this softshell 
turtle are comparable to what other authors have 
described about A. spinifera nesting habits. We were 
impressed by the last part of the digging strokes, but 
Minton (1972) also reported that softshells were vig- 
orously throwing the sand several meters away. 
Newman (1906) reported that nesting process usual- 
ly lasts about one hour and 535 completed the entire 
process in 44 minutes. Breckenridge (1960) reports 
almost exactly the same nesting observations from 
Minnesota as ours, in terms of date, time, duration 
and behaviour. According to Newman (1906) nest- 
ing can also occur at midday. 

WTiat was most notable about our observations was 
the daily pattern of activity preceding nesting. The 
day seemed divided into 3 parts. Morning would be 
spent basking and moving around in one area. Around 
noon, the turtle would leave the area and spend the 
whole afternoon swimming vigorously. By the end of 
the afternoon, the turtle would stop and then seem to 
be looking around for a suitable nesting spot. 

Our observations raised the following questions: 
Did the turtle choose this specific nesting site 
because it did not find any other suitable site while 
travelling or was it going for this exact spot because 
it knew it from past nesting experiences or because 
she had hatched there? We know that 535 was in this 
part of the river during the 1997 nesting season but it 
was not observed in nesting behaviour. What was the 
influence of the area's other nests in 535 choice? 
Smell seemed to have played a significant role in her 
search. We believe that this is what brought her out 
of the water the day it nested. She first moved on 
land at the beginning of a sharp curve probably 
smelling the nests at the other end of this curve, only 
a few meter away through forest. As reported by 
Fletcher (1996*), our observations indicated that 
Apalone spinifera can move considerable distance in 
a short period of time, exploring different potential 
nesting sites. This would suggest that confirmation 



2002 



Daigle, Galois, and Chagnon: Nesting of Softshell 



107 



of the nesting site of any specific turtle can only be 
achieved through direct observation of the laying 
process. 

Acknowledgment 

We thank Roger Bider, Joel Bonin, Lyne 
Bouthillier, Jocelyne Brisebois, Clement Lanthier, 
Martin Leveille, Jacques Jutras, David Rodrigue and 
Louis-Marc Soyez who provided help in the capture 
and tracking of the softshells during the whole pro- 
ject. David Rodrigue made useful comments on the 
first draft of this manuscript. Funding for the project 
was provided by the Ministere de I'Environnement 
et de la Faune du Quebec, Fondation de la faune du 
Quebec, Plan d' Action Saint-Laurent, and Societe 
d'Histoire Naturelle de la Vallee du Saint-Laurent. 

Document Cited (marqued * in text) 

Fletcher, M. 1996. Management of softshell turtle habi- 
tat, year 1, 1996. Upper Thames river Conservation 
Authority, London, Ontario. 23 pages. 

Galois, P. 1999. Recherche de sites de nidification de la 
tortue-molle a epines (Apalone spinifera) a la riviere aux 
Brochets et inventaires de I'espece sur la riviere Riche- 
lieu et la riviere des Outaouais. Gouvemement du Que- 



bec, Faune et Pares, Service de I'amenagement et de 
I'exploitation de la faune, Longueuil. Plan d' intervention 
sur la tortue-molle a epines au Quebec, rapport d'etape 
1998, xii + 95 pages. 

Literature Cited 

Breckenridge, W. J. 1960. A Spiny Soft-shelled Turtle 
nest study. Herpetologica 16: 284-285. 

Eigenman, C. H. 1896. Testudinae of Turkey Lake, 
Indiana. Proceedings of the Indiana academy of science. 
1895,5:262-264. 

Ernst, C. H., J. E. Lovich, and R. W. Barbour. 1994. 
Turtles of the United States and Canada. Smithsonian 
Institution Press, Washington and London. 578 pages. 

Evermann, B. W., and H. W. Clark. 1920. Lake Maxi- 
nkuckee, a physical and biological survey. The Depart- 
ment of Conservation, State of Indiana. Volume 1, 660 
pages. 

Minton, S. A. 1972. Amphibians and reptiles of Indiana. 
Indiana Academy of Sciences Monograph 3 : 1-346. 

Newman, H. H. 1906. The habits of certain tortoises. The 
Journal of Comparative Neurology and Psychology 16 : 
126-152. 



Received 25 April 2000 
Accepted 28 January 2002 



i 



Long-distance Movements by Female White-footed Mice, 
Pewmyscus leucopus, in Extensive Mixed-wood Forest 



Thomas J. Maier 

Northeastern Research Station, USDA Forest Service, Holdsworth Natural Resources Center, University of Massachusetts, 
Amherst, Massachusetts 01003-9285 USA 

Maier, Thomas J. 2002. Long-distance movements by female White-footed Mice, Pewmyscus leucopus, in extensive 
mixed-wood forest. Canadian Field-Naturalist 116(1): 108-1 11. 

Two adult female White-footed Mice {Pewmyscus leucopus) were recovered 14 730 m and 6840 m from where they were 
originally captured and tagged in central Massachusetts. Similar combinations of factors throughout an extensive forested 
landscape, including poor acorn {Quercus spp.) crops, high population densities of mice, and the exclusive social behaviour 
of other female mice, may have been responsible for these long distance movements. 

Key Words: White-footed Mouse, Pewmyscus leucopus, dispersal, distance, mixed- wood forest, movement, Quercus, 
small mammals, social dynamics, Massachusetts. 



Movement in space-time remains a little under- 
stood process for most organisms (Turchin 1998). 
Even less understood are long-distance movements 
(Koenig et al. 1996), such as those made by small 
mammals dispersing across landscapes (Zollner and 
Lima 1999; Andrzejewski et al. 2000; Bowman et al. 
2001). Nonetheless, even sporadic long-distance 
movements by small mammals may profoundly 
affect species' evolution, population dynamics, and 
community structure (Cockburn 1992; Allen et al. 
1993; Berlow 1999). As such, recognition of the spa- 
tial scale of such movements and their proximal 
causes becomes integral to understanding certain 
wildlife requirements, the effects of habitat perturba- 
tion, and possible epidemiological risks (Dickman et 
al. 1995; Kozakiewicz and Szacki 1995; Calisher et 
al. 1999). Here I report long-distance movement 
events by two female White-footed Mice (Pero- 
myscus leucopus) in extensive, New England mixed- 
wood forest, and discuss these movements in relation 
to the associated environmental conditions. 

Observations 

One month after capture on a small mammal trap- 
ping grid within the Quabbin watershed in central 
Massachusetts (42''27'13.7"N, 72°21'03.7"W), a 
tagged White-footed Mouse was recaptured 14 730 
m north of the original capture site. The adult female 
(21.2 g), apparently healthy and not exhibiting any 
signs of reproductive activity, was ear-tagged with a 
No. 1 Monel tag (National Band and Tag Co., 
Newport, Kentucky, USA) on initial capture, 15 
October 1998, and subsequently recaptured twice 
more at the same site. Approximately 4 weeks later, 
this mouse was trapped within a residence. Its identi- 
ty was verified based upon gender and the fact that 
no other small mammal researchers working in the 
area were using tag numbers in this sequence; a 



necropsy was not performed. A second tagged 
White-footed Mouse was trapped at another resi- 
dence 750 m northeast of the same Quabbin trapping 
grid during the same time period. This mouse was 
disposed of before verification of its identity. In mid- 
April 1993, the desiccated remains of another tagged 
White-footed Mouse were found outside a third resi- 
dence 6840 m north of its original capture site. This 
adult female, when captured on another small mam- 
mal trapping grid within the Quabbin watershed 
(42°26'50.9"N, 72°19'29.8"W) on 12 August 1992 
(its only capture), appeared healthy and did not 
exhibit any signs of reproductive activity. Its identity 
was verified as described above. 

Both trapping grids on which these mice were ini- 
tially captured were located within a fenced portion of 
the Quabbin watershed, with vehicle entry restricted 
by locked gates (this property is managed by the 
Metropolitan District Commission to provide water 
for the Boston municipal area). The landscape 
between the initial capture and final recovery sites 
consisted of contiguous, closed-canopy mixed-wood 
forest of a predominately Red Oak {Quercus rubra)- 
White Pine {Pinus strobus)-R.ed. Maple {Acer rubrum) 
forest cover type (Eyre 1980) with scattered residen- 
tial and agricultural openings. For the two mice travel- 
ing the furthest, potential travel routes were bisected 
by a minimum of six light-duty forest roads, three 
small streams, a two-lane primary highway, and a 
powerline right-of-way. Geomorphological features 
run mostly north-south and range in elevation from 
200-380 m. Distances, unless otherwise specified, 
were measured using a Rockwell PLGR-i-96 Global 
Positioning System receiver with a minimum horizon- 
tal accuracy of 10 m, and analyzed using ARC/INFO 
on a Sun SPARC station. 

Other environmental conditions characterizing the 
1998 period included high White-footed Mouse den- 



108 



2002 



Maier: Long-distance Movements by Mice 



109 



sity at the Quabbin trapping site (136/ha) and adja- 
cent sites (58-89/ha) (T. Maier, unpublished data). 
Few mice were actively breeding at this time, but 
large numbers of juveniles were observed (T. Maier, 
unpublished data). In comparison, historic densities 
for these mice for similar local habitat at the same 
time of year have ranged from 1 to 126/ha, with a 
mean density of 27.8/ha, SD = 30.6 (9 years data; W. 
Healy, USDA Forest Service, personal communica- 
tion). Area acorn {Quercus spp.) crops yielded > 75% 
less than the previous year, with virtually the entire 
1998 crop consumed by Acorn Weevil (Curculio 
spp.) larvae (R. Field, USGS Biological Resources, 
personal communication; W. Healy, personal com- 
munication). Local cUmatological statistics for a 44- 
day period in 1998 encompassing the potential travel 
of mice were a mean maximum temperature of 10.5° 
C; mean minimum temperature of -0.9° C; mean tem- 
perature of 4.9° C, and total precipitation of 24.3 mm, 
all as rain from five precipitation events (NWS*). 
Environmental conditions characterizing the 1992 
period beginning in August similarly included high 
White-footed Mouse density at the Quabbin trapping 
site (81/ha) and adjacent sites (40- 126/ha), and the 
absence of an acorn crop throughout the area, with 
mice populations dramatically declining by October 
of the same year (Elkinton et al. 1996). All popula- 
tion density estimates were generated using program 
CAPTURE (Otis et al. 1978; White et al. 1982). 

Discussion 

Although the distance covered by the furthest trav- 
eling White-footed Mouse reported here supersedes 
previous reports of long-distance movement by either 
homing or non-homing individuals of this species 
(2 km, Wegner and Henein 1991), the genus 
Peromyscus (3.2 km, Murie and Murie 1931), and the 
family Muridae (14 km, Dickman et al. 1995), travel 
over the 14.7 km tract during the period from initial 
to final capture was well within the potential capabili- 
ty of Peromyscus mice. Murie and Murie (1931) 
reported a Deer Mouse {Peromyscus maniculatus) 
returning over 3 km to its capture site in 2 days. 
Similarly, Calisher et al. (1999) reported an adult 
Deer Mouse homing 1200 m within 24 hrs. Dice and 
Hoslett ( 1 950), comparing the performances of seven 
species of Peromyscus on activity wheels, reported a 
maximum 24-hr record the equivalent of 37 km of 
travel by an individual White-footed Mouse. Besides 
possessing the stamina to travel great distances in 
short amounts of time, these mice have additionally 
been observed to surmount various obstacles to travel 
(Sheppe 1965; Teferi and Millar 1993; Calisher et al. 
1999) similar to those found in this observation. The 
possibility that these mice may have been transported 
via human conveyance cannot be totally discounted, 
but this seems unlikely given that the mice were ini 
tially captured within a restricted area behind locked 



gates. Additionally, personal interviews with the resi- 
dents who collected these mice revealed that their 
closest physical proximity consisted of one or two 
instances of a briefly parked vehicle (< 4 h during 
dayhght) approximately 2.5 km from where the 1998 
mice were initially captured. 

Most dispersal {sensu Lidicker and Stenseth 1992) 
by small rodent species has been observed to occur 
over short distances (McShea and Madison 1992; but 
see Koenig et al. 1996), with longest distances associ- 
ated with travel by males for most species. What fac- 
tors may have contributed to long-distance travel by 
female White-footed Mice? First, the contiguous for- 
est was likely advantageous to long-distance move- 
ments, compared to a more fragmented landscape 
(Yahner 1983; Krohne and Hoch 1999). Second, food 
deprivation has resulted in increased activity by Pero- 
myscus (Falls 1968 and references therein). Although 
Acorn Weevil larvae infestations may represent a sig- 
nificant dietary supplement, they are unsuitable as a 
primary energy source compared to sound acorns 
(Semel and Andersen 1988; Steele et al. 1996) from 
autumn through spring (Kirkpatrick and Pekins 2002). 
Third, an increase in population density has been 
observed to increase the number of dispersing 
Peromyscus mice (Fairbairn 1978; Krohne et al. 
1984). Fourth, resident P. leucopus females may espe- 
cially exclude conspecific female immigrants when 
resources decline (Metzgar 1971; Wolff 1989). 
Together, these factors suggest that the combination 
of high mouse density and low food supply through- 
out a large extensively-forested area may have created 
a social environment that inhibited local immigration, 
as posited by Stickel (1968) and Anderson (1989), 
thereby increasing the probability of long-distance 
travel by emigrating female mice. Other reports of 
long-distance movement by small mammals through 
large areas of continuous habitat have suggested high 
population densities as the incentive (Clark et al.l988; 
Bowman et al. 1999). Nevertheless, as observed by 
Fairbairn (1978), such movement is not likely a sim- 
ple funcfion of density or its fluctuation, but rather 
relates to individual behaviour within a social struc- 
ture along with changing resource patterns. 

Proximal factors leading to long-range movements 
are difficult to elucidate; this may be due to the inter- 
action of contributing factors, and perhaps, in part, 
because so few observations of such movements exist. 
Various studies have attempted to assess the spatial 
mobility of small mammals using arrays of traps 
(Krohne et al. 1984; Liro and Szacki 1994; Dickman 
et al. 1995; Andrzejewski et al. 2000; Bowman et al. 
2001); yet, due to logistical and methodological con- 
straints, the probability of detection o{ marked ani- 
mals may be expected to decline exponentially with 
distance, resulting in the significant underestimation 
of long-distance movements (Koenig et al. 1996; 
Turchin 1998). The long-distance movements report- 



110 



The Canadian Field-Naturalist^ 



Vol. 116 



ed here may eventually prove more common than pre- 
viously thought, requiring those studies modeling the 
spatial distribution of similar small mammals to adopt 
a new metric. 

Acknowledgments 

I particularly thank T. Cox, M. Delano, D. Small, 
and J. Starkey for reporting the collection of these 
mice. I also thank D. Clark for site information; D. 
Goodv^in for analyzing distance data; R. Field and 
W. Healy for data on local small mammal density 
and acorn crops; G. Boettner, J. Bowe, Jr., R. Brooks, 
R. DeGraaf, K. Doyle, J. Hestbeck, D. King, J. 
Wolff, and P. Zollner for suggestions regarding the 
manuscript; S. Cooter for ethological instruction; and 
the Massachusetts Metropolitan District Commission, 
Division of Watershed Management, Quabbin 
Section, for access to their management areas. 

Documents Cited (marked with* in text) 

[NWS] National Weather Service, NOAA. Orange and 

Worcester, Massachusetts, USA. Available from: URL: 

http://tgsv.nws.noaa.gov/er/box/climate/ 

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Berlow, E. L. 1999. Strong effects of weak interactions in 
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Bowman, J. C, M. Edwards, L. S. Sheppard, and G. F. 
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Bowman, J., G. F. Forbes, and T. G. Diiwortli. 2001. 
Distances moved by small woodland rodents within 
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64-67. 

Calislier, C. H., W. P. Sweeney, J. J. Root, and B. J. 
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Clark, B. K., D. W. Kaufman, G. A. Kaufman, E. J. 
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Dice, L. R., and S. A. Hoslett. 1950. Variation in the 
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Dickman, C. R., M. Predavec, and F. J. Downey. 1995. 
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Elkinton, J. S., W. M. Healy, J. P. Buonaccorsi, G. H. 
Boettner, A. M. Hazzard, H. R. Smitli, and A. M. 
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Eyre, F. H. 1980. Forest cover types of the United States 
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Fairbairn, D. J. 1978. Dispersal of deer mice, Pero- 
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Falls, J. B. 1968. Activity. Pages 543-570 in Biology of 
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Koenig, W. D., D. Van Vuren, and P. N. Hooge. 1996. 
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Kozakiewicz, M., and J. Szacki. 1995. Movements of 
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Krohne, D. T., B. A. Dubbs, and R. Baccus. 1984. An 
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Krohne, D. T., and G. A. Hoch. 1999. Demography of 
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Lidkicker, W. Z., Jr., and N. C. Stenseth. 1992. To dis- 
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Sheppe, W. 1965. Dispersal by swimming in Peromyscus 
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Turchin, P. 1998. Quantitative analysis of movement: 
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Yahner, R. H. 1983. Population dynamics of small mam- 
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1019-1030. 

Received 4 July 2000 
Accepted 12 March 2002 



I 



Recent Trends in Stem Numbers in Goldenseal, Hydrastis canadensis. 
Populations at the Northern Limit of its Range 

Adrianne Sinclair^ and Paul M. Catling^ 

'Box 214 Metcalfe, Ontario KOA 2P0 Canada; and Biology Department, University of Ottawa, Ottawa, Ontario KIN 6N5 

Canada 
^Eastern Cereal and Oilseed Research Centre, Biological Resources Program, Agriculture and Agri-food Canada, Research 

Branch, William Saunders Building, Central Experimental Farm, Ottawa, Ontario KIA 0C6 Canada 

Sinclair, Adrianne, and Paul M. Catling. 2002. Recent trends in stem numbers in Goldenseal, Hydrastis canadensis, popu- 
lations at the northern limit of its range. Canadian Field-Naturalist 1 16(1): 1 12-1 15. 

A 15% increase in stem number occurred in 14 natural Ontario Goldenseal populations surveyed between 1998 and 2000, 
based on maps of stems in lm= quadrats. Increase occurred in 10 of the 14 populations and was significant. The increase 
mostly involved small stems, but neither seedlings nor very small plants were observed suggesting that stem increase was 
largely due to vegetative reproduction by rhizomes and/or root tips. These data correspond to expectation since populations 
have not expanded rapidly nor spread substantially over the past few decades, but rather appear to have expanded slightly 
and slowly. Corresponding to earlier speculation, evidence is accumulating to suggest that substantial spreading and large 
population increases may be dependent on specific kinds and levels of disturbance some of which are no longer prevalent 
or frequent. 

Key Words: Goldenseal, Hydrastis canadensis, rare plant, medicinal plant, population trends, demography, population 
growth rate, population structure, management, monitoring, southwestern Ontario 



The top-selling and increasingly popular (Small 
and Catling 1999; Foster 2000*) deciduous wood- 
land plant, Goldenseal {Hydrastis canadensis L.) 
was recently re-evaluated as threatened in Canada 
(COSEWIC 2000*) due to few scattered popu- 
lations, habitat loss, and potential threat of over- 
collection for the medicinal plant market (White 
1991*; Sinclair and Catling 2000a). The 20 popula- 
tions in Canada occur in southwestern Ontario at 
the northern limit of its native range, which extends 
from southern Wisconsin to southern Vermont and 
south to Arkansas and northern Georgia (Sinclair 
and Catling 2000a). The perennial rhizomes of 
Goldenseal, which produce an annual stem each 
year and enable vegetative reproduction, are col- 
lected for the medicinal plant market. The market 
demand for Goldenseal has been largely met by 
harvest of wild populations in the United States 
(Foster 1991; Small and Catling 1999; USFWS 
1997*), which are now considered endangered, crit- 
ically imperilled, imperilled, rare, or uncommon, 
depending on the State (USFWS 1997*). Apart 
from anecdotal observations, information on 
growth trends in Goldenseal populations is lacking 
for all parts of its range. Such information is essen- 
tial for the development of effective conservation 
strategies and monitoring programs, and the need 
for it has been emphasized in the most recent 
review of goldenseal ecology and population biolo- 
gy (Gagnon 1999*). Here we present trends in pop- 
ulation structure and growth rate, over two years, 
from 14 natural Goldenseal populations in south- 
western Ontario. 



Methods 

In May of 1998, a Im^ quadrat was placed at the 
edge of a single colony within each of 14 naturally 
occurring Goldenseal populations. The 14 popula- 
tions occur in Lake Erie Lowland Ecoregion 135 and 
the southwestern portion of adjacent Manitoulin- 
Lake Simcoe Ecoregion 134 of the Mixedwoods 
Plains Ecozone (Ecological Stratification Working 
Group 1995). Site numbers in Figures 2, 3, and 4 
correspond to locations (Sinclair and Catling 2000a) 
on file with the Committee on the Status of 
Endangered Wildlife in Canada (COSEWIC) [see 
Sinclair and Catling 2000a and references therein]. 
Each quadrat was permanently marked with four alu- 
minum pins and a numbered aluminum tag. All stems 
were mapped within each quadrat according to the 
following size classes: small (single leaf < 5 inches in 
width); large (single leaf > 5 inches in width); and 
flowering (two or more leaves). The flowering stems 
are produced by larger rhizomes which are presumed 
to be the oldest. The smallest stems are generally 
associated with small rhizomes or root tips. Presence 
or absence of seedlings was noted. Due to the clonal 
nature of the plant, all of the stems may have devel- 
oped from a single individual. The distinction 
between ramets and genets was beyond the scope of 
the present survey. In May of 2000, the 14 quadrats 
were revisited and remapped according to the same 
size classes. In July of both years, quadrats were 
revisited to record fruiting stems. The quadrat maps 
developed in 2000 were compared to those developed 
in 1 998 to document any change in stem numbers and 
size classes. Total number of stems, as well as 



112 



2002 



Sinclair and Catling: Stem Numbers in Goldenseal 



113 



700 




B F L 
Stem Class 

Figure L Histogram showing overall trend in population 
structure, from 1998 to 2000, based on a m- quadrat 
sample from each of 14 goldenseal populations in 
Ontario. B = fruiting, F = flowering, L = large non- 
flowering, LL = small non-flowering. 



numbers of small, large, flowering, and fruiting 
stems, over all sites, were compared between the two 
years using simple t-tests. 

Results and Discussion 

There was no evidence of harvest, or other recent 
disturbances (tree removal, trampling, etc.), at any of 
the 14 populations. Overall (14 quadrat samples com- 
bined), between 1998 and 2000, there was a signifi- 
cant increase of 15% (p = 0.0036, 550 to 635 stems. 
Figure 1). Ten out of 14 patches show an increase in 
the number of stems (Figure 2). The greater number 
of stems in 2000 is due to a significant increase in the 
number of small stems (p = 0.003, Figures 1 and 3). 
Almost all patches show an increase in small stems, 
and the increases are much greater than the losses 
(Figure 3). No seedlings or very small plants (< 1 
inch leaf width) were noticed, suggesting that stems 
developed from roots or rhizomes of existing plants 
and that effective recruitment by seed was negligible. 
These data suggest that patches are slowly expanding 
by vegetative reproduction. Fruiting stems have also 
increased with marginal significance (p = 0.045, 
Figures 1 and 4). Only two patches show a decrease 
in the fruiting component (Figure 4). The increase in 
fruiting stems was not associated with increased flow- 
er production: virtually no change occurred in flower- 
ing stems (Figure 1 ). The increased number of fruit- 
ing stems in 2000 is therefore a result of a greater 
proportion of flowering stems producing fruit, which 
may have been a consequence of the weather. 
Although reduced herbivory could also be an expla- 
nation, there was no evidence for this. No increase in 



o 

c 





</i 

CD 

0) 

1_ 

O 



Q 




1A 1B 2 3 5 6 7 11 121415 171820 
Site Number 

Figure 2. Increases or decreases in stem number, from 
1998 to 2000, based on a m^ quadrat sample from 
each of 14 goldenseal populations in Ontario. 



seed production capability actually occurred. Large 
stems have decreased but the loss is not statistically 
significant (Figure 1). 

These data correspond to expectation since popula- 
tions have not expanded rapidly nor spread substan- 
tially over the past few decades, but rather appear to 
have expanded slightly and slowly (Sinclair and 
Catling 2000a). The 15% increase in overall stem 
production and the increased fruit production were 



24 



(D 
(A 

O 

c 



(D 
(/) 
OJ 
0) 

o 

0) 

Q 



16- 



8- 







\ \ \ \ I I \ \ \ ^^ ^ I I 
-• • - 

- • 

•____•_ 



1A IB 2 3 5 6 7 11 12 141517 
Site Number 

Figure 3. Increases or decreases in number of sm; 
flowering stems, from 1998 to 2()()(). based 
quadrat sample from each of 14 Goldenseal 
tions in Ontario. 



18 20 

ill. non- 
on a m- 
popula- 



114 



The Canadian Field-Naturalist^ 



Vol. 116 



U) 
03 
0) 

O 

c 



0) 

(/) 

CD 
<D 

o 

Q 




1112 1415171820 



Site Number 

Figure 4. Increases or decreases in number of fruiting 
stems, from 1998 to 2000, based on a m^ quadrat 
sample from each of 14 Goldenseal populations in 
Ontario. 



possibly a result of the relatively warmer summers of 
1998 and 1999, which are characteristic of the main 
Goldenseal range further south. 

The lack of evidence for reproduction by seed sug- 
gests that a limitation must be operating on sexual 
reproduction other than pollination and dispersal (see 
Sinclair et al. 2000). The stability of Ontario 
Goldenseal patches correspond to trends in other 
woodland perennials, such as American Ginseng 
(Panax quinquefolius L.) and Wild Leek {Allium tric- 
occum Ait.), which are slow-growing with population 
growth rates near equilibrium (population stability or 
maintenance, Charron and Gagnon 1991; Nault and 
Gagnon 1993). Conversely, in disturbed habitats, 
where more resources such as light and nutrients are 
made available, plants tend to have highly variable 
population growth rates which can reach well above 
equilibrium (e.g.. Furbish' s Lousewort {Pedicularis 
furhishiae S. Wats.) and Downy Oatgrass (Danthonia 
sericea Nutt.), as noted by (Gagnon 1999). Since 
occurrences of Goldenseal have been associated with 
disturbance (Sinclair and Catling 2000b), it seems 
probable that substantial spread and increase in 
Goldenseal populations is disturbance-dependent. The 
results of recently initiated field experiments, along 
with further population monitoring, will enable a bet- 
ter understanding of disturbance effects and provide 
useful information for recovery and management. 

Acknowledgments 

Fieidwork was supported by the Endangered 
Species Recovery Fund (ESRF) of World Wildlife 
Fund Canada and the Canadian Wildlife Service of 



Environment Canada, and also by Agriculture and 
Agri-food Canada and the University of Ottawa. 
Keiko Lui, Josh Flanagan, and Bruce Sinclair provid- 
ed field assistance. Conservation Authorities and pri- 
vate landowners kindly allowed us to work on their 
properties. 

Documents Cited (marked * in text) 

COSEWIC (Committee on the Status of Endangered 
Wildlife in Canada). 2000. Canadian species at risk. 
May 2000. 28 pages, http:// www.cosewic.gc.ca/ 
COSEWIC/2000_list.pdf 

Foster, S. 2000. Goldenseal, Hydrastis canadensis: 
goldenseal's future. Steven Foster Group. http://www. 
stevenfoster.com/education/monograph/golden- 
seal.html. 

Gagnon, D. 1999. A review of the ecology and population 
biology of goldenseal, and protocols for monitoring its 
populations. Final report to the Office of Scientific 
Authority of the US Fish and Wildlife Service. Groupe 
de recherche en ecologie forestiere, Universite du Quebec 
a Montreal, http://www.nps.gov/plants/medicinal/pubs/ 
goldenseal. 

USFWS (United States Fish and Wildlife Service). 1997. 
Amendment to Appendix II of Convention on 
International Trade in Endangered Species of Wild Fauna 
and Flora (CITES), United States of America - Proposal 
to Include Hydrastis canadensis in Appendix II, in accor- 
dance with Article 2, paragraph 2A. 31 pages. 

White, D. J. 1991. Status report on the Golden Seal, 
Hydrastis canadensis, in Canada. Committee on the 
Status of Endangered Wildlife in Canada, Ottawa. 21 
pages. 

Literature Cited 

Charron, D., and D. Gagnon. 1991. The demography of 
northern populations of Panax quinquefolium (American 
ginseng). Journal of Ecology 79: 431^145. 

Ecological Stratification Working Group. 1995. A nation- 
al ecological framework for Canada. Agriculture and 
Agri-food Canada, Research Branch, Centre for Land and 
Biological Resources Research and Environment Canada, 
State of the Environment Directorate, Ecozone Analysis 
Branch, Ottawa/Hull. 18 pages + appendix + map. 

Foster, S. 1991. Go\dtr\sta\ Hydrastis canadensis. 
Botanical Series Number 309. American Botanical 
Council, Austin. 8 pages. 

Nault, A., and D. Gagnon. 1993. Ramet demography of 
Allium tricoccum, a spring ephemeral, perennial forest 
herb. Journal of Ecology 81: 101-119. 

Sinclair, A., and P. M. Catling. 2000a. Status of gold- 
enseal, Hydrastis canadensis (Ranunculaceae), in 
Canada. Canadian Field-Naturalist 114: 11 1-120. 

Sinclair, A. and P. M. Catling. 2000b. Ontario goldenseal 
populations in relation to habitat size, paths, and wood- 
land edges. Canadian Field-Naturalist 1 14(4): 652-655. 

Sinclair, A., P. M. Catling, and L. Dumouchel. 2000. 
Notes on the pollination and dispersal of goldenseal, 
Hydrastis canadensis L., in southwestern Ontario. 
Canadian Field-Naturalist 114: 499-501. 

Small, E., and P. M. Catling. 1999. Canadian medicinal 
crops. National Research Council of Canada, Ottawa. 
240 pages. 

Received 6 February 2001 
Accepted 12 March 2002 



2002 



Sinclair and Catling: Stem Numbers in Goldenseal 



115 



Addenda 

Corrections and additional notes for "Ontario 
Goldenseal, Hydrastis canadensis, populations 
in relation to habitat size, paths, and woodland 
edges". Canadian Field-Naturalist 114(4): 
652-655. 

A substantial interest in this earher article has lead to 
the following notes. One reader questioned whether or 
not paths in woodlands represent a random sample of 
the habitat? In other words paths may follow habitat 
subunits within woodlands, such as higher ground. 
Thus certain plant distributions in a forest may be cor- 
related with paths but without the paths themselves 
being a causal factor. The woodlands containing popu- 
lations of Goldenseal that we examined were either 
flat or uniformly and gradually hummocky. The paths, 
which were associated with the use of tractors or all- 
terrain vehicles, appeared to be a random sample of 
the habitat. The paths were not conspicuously associat- 
ed with openings in the canopy, although some of the 
edges certainly received more light than the interior. 
Consequently disturbance apparently benefitting 
Goldenseal did not appear to be light-dependent, 
although increased hght may contribute to a beneficial 
effect. 

Neither of the following two corrections changes the 
conclusions of the paper: 

(1) In calculating the effect of namral area size, all 
populations within a natural area should have been 
pooled. This applies to three cases (a) 11, 12, 13 and 



14, (b) 16 and 17, (c) lA and IB, all in table 1. 
Instead of "a marginally significant decrease in num- 
ber of Goldenseal stems with increasing natural area 
size", there was no significant relationship between 
number of Goldenseal stems and natural area size 
(P = 0.982, R- = 0.0). It is of interest to note here that 
when 3 large sites with large leverage were removed 
from the data set, there was still no significant rela- 
tionship with natural area size. 
(2) The figures in the D- value column in Table 1 are 
incorrect and so is the procedure described at the end 
of results in the first two sentences of the paragraph 
beginning "In order to ...". This should read "In 
order to ... disturbance values were assigned to each 
site based on a scale of 1-5 (1 = undisturbed, 5 = 
heavily disturbed). The assessment was based sub- 
jectively on a number of factors including gaps in 
the canopy, evidence of a cutting operation (bulldoz- 
ing, stumps, log dragging, etc.), trampling and vehi- 
cle tracks, and presence of invading alien species 
and/or native species of more open sites." Since this 
is actually what was done there is no change to the 
results or conclusions. The column of D values in 
Table 1 from top to bottom should be 3, 5, 2, 5, 1, 1, 
2, 1,2,5,3,2, 1, 1, 1,2,3, 1, l,and5. 

Adrianne Sinclair and Paul Catling 

Eastern Cereal and Oilseed Research Centre, Agriculture 
and Agri-food Canada, Research Branch, Wm. Saunders 
Building, Central Experimental Farm, Ottawa, Ontario 
KIA 0C6 Canada; e-mail: catlingp@em.agr.ca 



A Comparison of Techniques for Assessing Amphibian Assemblages 
on Streams in the Western Boreal Forest 



Cynthia A. Paszkowski\ Garry Scrimgeour^ Beverly A. Gingras', and Sharon Kendall^ 

'Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9 Canada; e-mail correspondence: 

cindy.paszkowski@ualberta.ca 
^Forest Resources Business Unit, Alberta Research Council, P.O. Bag 4000, Vegreville, Alberta T9C 1T4 Canada 

Paszkowski, Cynthia A.. Garry Scrimgeour, Beverly A. Gingras, and Sharon Kendall. 2002. A comparison of techniques 
for assessing amphibian assemblages on streams in the western boreal forest. Canadian Field-Naturalist 116(1): 
116-119. 

The western boreal forest of Canada is rapidly being altered by agriculture, forestry, and energy extraction. As part of the 
Alberta Forest Biodiversity Monitoring Program's effort to develop monitoring schemes for biota associated with streams, 
we compared the performance of four sampling techniques for amphibians (constrained visual searches, call surveys, pitfall 
traps, and above-ground funnel traps) in June through August along two low gradient streams. Of the four anuran amphib- 
ian species that occur in the region, we encountered adults and young-of-the-year of two. Wood Frog (Rana sylvatica) and 
Western Toad {Bufo boreas), with the former species dominating our surveys. Given the time and equipment constraints 
imposed by the monitoring-program protocol, visual surveys proved to be the most effective technique (88.5% of amphib- 
ian records) for simply determining the presence of species. Pitfall traps performed better than funnel traps. Call surveys 
were the least effective technique principally because sampling took place after most breeding was completed. 

Key Words: Alberta, biomonitoring, pitfall trap, visual survey, sampling techniques. Western Toad, Bufo boreas. Wood 
Frog, Rana sylvatica. 



Within the last decade the boreal forest in Alberta, 
which covers the northern half of the province 
(Rowe 1972), has become increasingly disturbed and 
fragmented by agricultural expansion, logging, ener- 
gy extraction, and road building. The cumulative 
effects of municipal and industrial activity on boreal 
ecosystem integrity and biodiversity are receiving 
increasing attention. The Alberta Forest Biodiversity 
Monitoring Program (AFBMP) was established in 
1996 to identify a suite of quantifiable biological 
attributes (i.e., variables ranging from the individual 
to the community-level) to serve as indicators of sus- 
tainable forest management and to develop sampling 
protocols for detecting changes in terrestrial and 
aquatic biodiversity within Alberta's 500 000 km^ 
forested land base (Schneiderl997*). 

AFBMP is based on a network of 1250 sites located 
on a systematic grid. Individual sites will be moni- 
tored once every 5 years with 250 sites visited annual- 
ly. Each site will include terrestrial habitat to be sam- 
pled for nonvascular and vascular plants, arthropods, 
birds, and mammals. The terrestrial habitat will be 
paired with a nearby stream location. Forest streams 
are sensitive to catchment disturbances and respond 
quickly to changes in adjacent habitats (Power et al. 
1988; Welsh and Ollivier 1998). Because sites are 
numerous and remote (crews will typically fly in), 
sampling of all stream biota (benthic algae, macroin- 
vertebrates, fish) and habitat features will be complet- 
ed by a two-person crew in 2 - 4 days with only 0.5 - 
1 .5 days allotted to any particular taxonomic group. 
Sampling will occur in June, July, and August. 



Because of the presumed sensitivity of many 
species to degradation of both terrestrial and aquatic 
habitats (Green 1997), amphibians would seem a 
natural group for inclusion in AFBMP surveys. 
Likewise, the ambitious scope of the program offers 
a unique opportunity to increase our meager knowl- 
edge of the distribution and ecology of the six anu- 
ran and two caudate amphibians that occur in 
Alberta's forests, particularly elucidating their use of 
streams and adjacent riparian habitat (Russell and 
Bauer 2000). However, given the time and resource 
constraints of the proposed survey design and well 
documented natural fluctuations in amphibian num- 
bers (Semlitsch 2000), two questions emerge: "What 
sampling techniques perform best?" and "Will sur- 
vey results support the use of amphibians as environ- 
mental indicators by AFBMP or other large-scale 
monitoring programs?' 

To address these questions, our study compared 
four techniques for sampling amphibians (con- 
strained visual searches, call surveys, pitfall traps, 
and above-ground funnel traps) along two low gradi- 
ent streams in the boreal forest of northeastern 
Alberta. Surveys adopted the previously described 
constraints on effort, seasonal timing, and equipment 
prescribed by AFBMP stream protocols. 

Materials and Methods 

In 1999 amphibian assemblages were sampled 
along two third-order forest streams. Bear Creek 
(55°44'55"N, 112°12'16"W) and Crow Creek 
(55°37'26"N, 112°ir36"W), north of Wandering 



116 



2002 



Paszkowski, Scrimgeour, Gingras, and Kendall: Comparison 



117 



River, Alberta. Sites were within the known ranges 
of four species of amphibians (all anurans) that are 
widespread in boreal Alberta (Russell and Bauer 
2000): Western Toad {Bufo boreas), Canadian Toad 
{Bufo hemiophrys). Boreal Chorus Frog. {Pseudachs 
maculata), and Wood Frog (Rana sylvatica). 

Visual surveys 

Amphibians were surveyed three times at each 
stream: June (14 - 20), July (12 - 16) and August (23 
- 27). During a survey period, the two-person team 
performed low intensity area-constrained visual 
searches (Crump and Scott 1994) on two dates 
between 08:00 and 12:00 MDST. Searches were 
conducted along 200 m long x 1 m wide flagged 
transects, one on each side of the stream at a distance 
of 1 - 5 m from the streambed. One person walked 
slowly searching for animals active on the surface. 
Animals were captured by hand, identified to 
species, measured (snout-urostyle length, SUL), 
weighed, aged based on size and date of capture 
("adults" = 1-i-year old, any individual captured in 
June or individuals > 27 mm SUL in July-August; 
young-of-the-year = newly metamorphosed individ- 
uals captured in July-August, < 27 mm SUL; based 
on C. Paszkowski, unpubhshed data for Wood Frog 
and Western Toad), then released. Forty-two percent 
of the individuals observed during surveys escaped 
capture. These animals were identified to species and 
assigned an age if possible. The location of each 
amphibian encountered (captured or observed) was 
recorded based on markers placed at 50-m intervals 
along the transect. The total time required to survey 
a transect ranged from 18 to 60 min. To control for 
effort, we calculated the total number of animals 
encountered per hour of searching. 

Call surveys 

At both sites, we performed call surveys with 
and without recording devices. Surveys with a "live" 
listener followed the audio strip technique 
(Zimmerman 1994). These were performed twice per 
month (one survey per date), at the same time and on 
the same 200 m transect as visual surveys. 

We used microcassette recorders (Panasonic RN- 
202, Matsushita Electric Industrial Co., Ltd.) for 
fixed-position call surveys at each stream, twice each 
month, between 08:00 and 12:00. A recorder was 
placed on each side of the stream inside a plastic jug 
located 1 .5 m up a tree. The two recorders were sep- 
arated by an additional 200 m measured linearly 
along the stream bank. Each recording session lasted 
60 min, resulting in 4 h of recording per site each 
month. 

Trapping 

In June a team of two workers installed four pitfall 
arrays and six funnel-trap arrays at the two sites. 
Arrays were equally divided between the two stream 
banks, 2 - 10 m from the water, and were separated 



by 50 m. Pitfall arrays consisted of four traps linked 
by drift fence, 30 cm high x 8 m long polyethylene 
strips attached to wooden stakes and anchored in the 
soil. Two traps were centrally located, on either side 
of the drift fence, and one trap was located at the end 
of either arm. Traps were buried plastic buckets, 24 
cm deep and 24 cm wide, fitted with 7 mm- wire 
mesh funnels with 8 cm openings. Damp sponges 
were placed in each trap and twigs were inserted ver- 
tically to allow small mammals to escape. Arrays 
were left intact between monthly sessions, but traps 
were closed with plastic lids. 

Funnel-trap arrays consisted of two collapsible 
fiber-mesh minnow traps (50 cm long x 25 cm high 
X 25 cm wide with two inward directed openings 
5.5 cm in diameter; Nylon Net, Memphis, Tennesse) 
centrally located on either side of a drift fence (as 
described above). Litter was used to partially cover 
traps and to create "ramps" to funnel openings. 
Damp sponges were placed inside. Funnel traps were 
removed between monthly sessions, but drift fences 
were left intact. 

During the three monthly sampling sessions, pit- 
fall and funnel traps were open for 3 - 6 consecutive 
days and checked daily before noon. Information on 
captured amphibians was recorded as described for 
searches. For pitfall and funnel traps, we calculated 
the number of animals caught per trap-night ( 1 trap- 
night = 1 trap opened for 24 h). 

Results 

Visual surveys were the most effective sampling 
technique, in terms of the combined number of cap- 
tures and observations (115), followed by pitfall 
arrays (10), funnel arrays (3) and live call surveys 
(2). Automated call surveys failed to record any 
amphibians. The two males heard calling were 
Western Toads at Crow Creek in June. Two Western 
Toads, one adult (37 mm SUL, pitfall trap) and one 
young-of-the-year (19 mm SUL, visual survey), 
were also captured at Crow Creek. All other amphib- 
ians encountered were Wood Frogs. The size distri- 
bution of Wood Frogs for all sampling combined 
was similar at the two sites: Bear Creek: mean SUL 
(mm) = 31.0 ± 9.4 (SD), range: 20 - 52.5, N = 35; 
Crow Creek: mean SUL = 39.8 ± 8.6, range: 20 - 
59.5; N = 42. Most frogs captured were adults but 
we did record young-of-the-year (N = 19; 20 - 27 
mm SUL) at both sites in July and August. 

Survey results differed between sites and among 
months. Encounter rates during visual surveys and 
capture rates with funnel traps were greater at Crow 
Creek (Figure la, b). Pitfall capture rates were 
greater at Bear Creek (Figure lb). Visual survey 
encounter rates and funnel-trap capture rates gradu- 
ally decreased as the summer progressed (Figure Ic, 
d), whereas pitfall capture rates increased in August, 
reflecting captures of young-of-the year (Figure Id). 



118 



The Canadian Field-Naturalist. 



Vol. 116 




0.050 



S* 0.040 - 



a. 
n 



« 0.030 
o 



£ 0.020 

3 
Q. 

s 

•g 0.010 

6 

z 

0.000 



M 




Bear 



Crow 



(A 

'E 
a. 
2 



ra 0.030 



"5 0.010 



0.000 



■ visual survey 
□ pitfall traps 
gj funnel traps 




June 



July 



August 



Figure 1 . Encounter and capture rates for the Wood Frog at Bear Creek and Crow Creek using visual surveys (a), and pit- 
fall and funnel traps (b) for the three monthly sampling sessions combined. Monthly encounter and capture rates for 
the Wood Frog using visual surveys (c), and pitfall and funnel traps (d) for the two sites combined. Total time 
devoted to visual surveys was 8.05 h at Bear Creek and 8.12 h at Crow Creek. Trapping effort totaled 216 pitfall 
and 156 funnel trap-nights at Bear Creek, and 188 pitfall and 140 funnel trap-nights at Crow Creek. 



Discussion 

We documented substantial differences in the per- 
formance of the four sampling techniques. Live and 
automated call surveys largely failed. The proposed 
AFBMP schedule for sampling streams simply does 
not coincide with the breeding season of anuran 
amphibians in boreal Alberta. Calling begins in late 
April, peaks in May and only Bufo species (which 
we did detect) continue breeding into June (Russell 
and Bauer 2000). Although relatively quick and 
easy, call surveys make little biological sense in this 
program. 

Pitfall and funnel trapping sessions resulted in few 
amphibian records, accounting for only 10% of the 
total number of individuals. This was a disappointing 
performance considering that the two workers 
required 18 h and 13.5 h to install the full suite of 
pitfall and funnel-trap arrays, respectively. Using our 
pitfall trapping effort and design (16 traps in 4 
arrays), two overnight trapping sessions were 
required to yield a 70% probability of capturing one 
Wood Frog. Funnel traps never even reached this 
level of performance. Trapping offers a reliable tech- 



nique at sites where low travel costs and easy access 
permit sampling over an extended time period, par- 
ticularly in upland habitats where amphibian densi- 
ties are relatively low or occurrence sporadic (Szaro 
et al. 1988). Pitfall trapping did result in the capture 
of newly metamorphosed Wood Frogs, thus provid- 
ing evidence of local breeding, possibly in Beaver 
(Castor canadensis) impoundments which were 
common on both streams. 

Visual searches required relatively small invest- 
ments in time and materials and proved to be the 
most successful technique for sampling streams. 
Wood Frogs were seen on 22 of 24 surveys. During 
these 22 surveys, searching along a transect for only 
50 m offered a 70% probability of encountering at 
least one Wood Frog. Like pitfall trapping, visual 
searches encountered young-of-the year frogs. 
Visual surveys were well suited to the study streams 
because riparian areas were characterized by a low 
occurrence of woody debris and relatively sparse 
understory, features not shared by all forest streams 
in Alberta. Even our searches yielded fewer amphib- 
ians as the summer progressed and streamside grass- 



4 



2002 



Paszkowski, Scrimgeour, Gingras, and Kendall: Comparison 



119 



es and forbs increased in height and density. Thus, 
although constrained visual searches might best meet 
AFBMP criteria, data generated by a single visit to a 
location will be strongly influenced by habitat struc- 
ture, season, weather, time of day, and surveyor 
competence (Crump and Scott 1994). 

Our surveys encountered only two of four anuran 
amphibians that occur in northeastern Alberta 
(Russell and Bauer 2000). As Roberts and Lewin 
(1979) reported from visual surveys conducted 20 
years earlier, the Wood Frog overwhelmingly domi- 
nated our records. The absence of the common, but 
cryptic. Boreal Chorus Frog can be attributed to 
sampling biases linked to season (nonbreeding ver- 
sus breeding and calling) and habitat (streams versus 
ponds). The paucity of toads may reflect insufficient 
sampling effort to detect these less abundant and 
more fossorial species. 

If AFMBP maintains its proposed sampling proto- 
cols and adopts constrained visual searches to moni- 
tor forest amphibians, our study suggests that the 
program will reliably document the presence/ 
absence of the Wood Frog at sites and collect some 
demographic information on poorly known western 
and northern populations (e.g., Leclair et al. 2000). It 
seems less likely that the program will be able to 
assess the health of boreal forest populations of 
amphibian species like the Canadian Toad which are 
of conservation concern in Alberta (Hamilton et al. 
1998). In closing, we support amphibian surveys as 
part of innovative, large-scale projects such as the 
AFBMP. However, programs designed to sample 
simultaneously a large number of taxa over a large 
number of sites will inevitably result in compromises 
in terms of the types of data that can be collected for 
any given group of organisms and the accuracy of 
information concerning the distribution and status of 
species of interest. 

Acknowledgments 

We thank F. R. Cook, R. Seigel, and three anony- 
mous reviewers for providing comments on earlier 
drafts of this manuscript, and S. Boss for help in 
manuscript preparation. 

Documents Cited 

Schneider, R. 1997. Ecological diversity monitoring 
framework. Draft Discussion Paper, Alberta Forest 
Biodiversity Working Group, Edmonton, Alberta. 
www.fmf.ab.ca/p2.html. 



Literature Cited 

Crump, M. L., and N. J. Scott. 1994. Visual Encounter 
Surveys. Pages 84-92 in Measuring And Monitoring 
Biological Diversity: Standard Methods For Amphibians. 
Edited by W. R. Heyer, M. A. Donnelly, R. W. 
McDiarmid, L. C. Hayek, and M. S. Foster. Smithsonian 
Instimte Press, Washington. D.C. 

Green, D. M. Editor. 1997. Amphibians in decline: 
Canadian studies of a global problem. Herpetological 
Conservation, Volume 1 . 

Hamilton, I. M., J. L. Skilnick, H. Troughton, A. P. 
Russell, and G. L. Powell. 1998. Status of the 
Canadian Toad {Bufo hemiophrys) in Alberta. Alberta 
Environmental Protection, Wildlife Management 
Division, and the Alberta Conservation Association, 
Wildlife Status Report Number 12. Edmonton, Alberta. 

Leclair, R. Jr., M. H. Leclair, J. Dubios, and J. Daoust. 
2000. Age and size of Wood Frogs, Rana sylvatica, 
from Kuujjuarapik, Northern Quebec. Canadian Field- 
Naturalist 114: 381-387. 

Power, M. E., R. J. Stout, C. E. Gushing, P. P., Harper, 
F. R. Hauser, W. J. Matthews, P. B. Moyle, B. 
Statzner, and L R Wais De Badgen. 1988. Biotic and 
abiotic communities. Journal of the North American 
Benthological Society 7: 1-25. 

Roberts, W. and V. Lewin. 1979. Habitat utilization and 
population densities of the amphibians of northeastern 
Alberta. Canadian Field-Naturalist 93: 144-154. 

Rowe, J. F. 1972. Forest Regions of Canada. Canadian 
Forest Service 1300, Department of the Environment, 
Ottawa. Ontario. 

Russell, A. P., and A. M. Bauer. 2000. The Amphibians of 
Alberta. Second Edition. University of Calgary Press, 
Calgary, Alberta. 

Semlitsch, R. D. 2000. Principles for management of 
aquatic breeding amphibians. Journal of Wildlife 
Management 64: 615-631. 

Szaro, R. C, K. E. Severson, and D. R. Patton. Editors. 
1988. Management of amphibians, reptiles and small 
mammals in North America. U.S. Department of 
Agriculture, Forest Service, General Technical Report 
RM-166. 

Welsh, H. H. Jr., and L. M Ollivier. 1998. Stream 
amphibians as indicators of ecosystem stress: a case 
study from California's redwoods. Ecological 
Applications 8: 1118-1132. 

Zimmerman, B. L. 1994. Audio strip transects. Pages 92- 
97 in Measuring And Monitoring Biological Diversity: 
Standard Methods For Amphibians. Edited by W. R. 
Heyer, M. A. Donnelly, R. W. McDiarmid. L. C. Hayek, 
and M. S. Foster. Smithsonian Institute Press, 
Washington. D.C. 

Received 16 March 2001 
Accepted 26 February 2002 



Notes 



A Rare Leucistic Spiny Dogfish, Squalus acanthias, from the Bay of 
Fundy, Nova Scotia 



Brian W. Coad^ and John Gilhen^ 

'Canadian Museum of Nature, P. O. Box 3443, Station D, Ottawa, Ontario KIP 6P4 Canada 

^Nova Scotia Museum of Natural History, 1747 Summer Street, Halifax, Nova Scotia B3H 3A6 Canada 

Coad, Brian W., and John Gilhen. 2002. A rare leucistic Spiny Dogfish, Squalus acanthias, from the Bay of Fundy, Nova 
Scotia. Canadian Field-Naturalist 116(1): 120-121. 

A rare leucistic specimen of the Spiny Dogfish, Squalus acanthias, is reported for the first time in Nova Scotia, Canada and 
North America. It is a 915 mm total length adult female bearing pups. This specimen is an overall yellowish colour but has 
a large, darkly pigmented area on the head dorsally, a spot at the first and second dorsal fin origins, and blotches on the 
caudal peduncle and in the pectoral axils. It was captured while dragging for flounder in 70 fathoms about 2 km off Boars 
Head, Bay of Fundy, Nova Scotia, Canada at 44° 23'N, 66° 15'W, on 22 May 1989 by Stewart Taylor, Captain of the long- 
liner Nadia C of Centreville, Digby County. 

Key Words: Spiny Dogfish, Squalus acanthias, leucism, first record, Bay of Fundy, Nova Scotia, Canada. 



On 22 May 1989, Stewait Taylor, Captain of the 
longliner Nadia C of Centreville, Digby County, was 
dragging for flounder in 70 fathoms, at a place 
known to fishermen as "Bear Cove Ground" about 
2 km off Boars Head, Bay of Fundy, Nova Scotia, 
Canada at 44° 23'N, 66° 15'W, when he also cap- 
tured a leucistic Spiny Dogfish, Squalis acanthias. It 
is a 915 mm total length adult female bearing pups, 
and is catalogued in the Nova Scotia Museum of 
Natural History as NSM85308. 

Spiny Dogfish are normally grey to black in over- 
all colour with conspicuous white spots on the flanks 
and a white belly. The Bay of Fundy specimen is an 
overall yellowish colour but has a large, darkly pig- 
mented area on the head dorsally, a spot at the first 
and second dorsal fin origins, and blotches on the 
caudal peduncle and in the pectoral axils (Figure 1). 

Albinism or leucism is rare in sharks, rays and 
their relatives although relatively common in bony 
fishes (Dawson 1964; 1966; 1971; Dawson and Heal 
1976). Ben Brahim (1998) lists only 21 cases for 18 
species in all the elasmobranchs. Additional records 
are given in Gopalan (1971) and Talent (1973) for a 
count of 24 cases for 20 species, 8 being total albi- 
nos, 12 being "partial" albinos (presumably leucis- 
tic), and 4 not clearly defined. Leucistic specimens 
have a paleness of colouration (Lincoln et al. 1982) 
with the eyes usually dark. The first record of a 
leucistic Spiny Dogfish in Canadian waters is there- 
fore of some rarity. 

Selected measurements on the specimen preserved 
in ethanol are as follows, with the methodology after 
Compagno (1984). Figures in parentheses are ranges 



for comparative material (NMC 66-600, 556 mm 
total length (TL), Nova Scotia, George's Bank, ca. 
42°00'N, 67°00'W; NMC 84-1512, 623 mm TL, off 
Sable Island, ca. 44°18'N, 59°45'W; NMC 68-19, 
665-704 mm TL, Newfoundland, St. John's, White's 
Ledge, ca. 47°42'N, 52°42'W; NMC 67-9, 80 mm 
TL, New Brunswick, Saint John River, 45°22'N, 
66°12'W). 

Prenarial length in preoral length 2.0 (1.8-2.0); 
narrowest mouth width in preoral length 1.5 
(1.3-1.7); preorbital length in prebranchial length 
2.6 (2.4-2.6); eye length in preorbital length 2.1 
(1.9-2.3); first dorsal fin spine length in first dorsal 
fin height 1.4 (1.1-1.5); first dorsal fin height in fin 
length 2.1 (1.9-2.1); the second dorsal fin margin in 
first dorsal fin margin 1.3 (1.2-1.3). These morpho- 
metric characters fall within the ranges for normally 
coloured Spiny Dogfish and the leucistic specimen 
agrees with the description in Compagno (1984) in 
regards to other characters. 

The specimen is leucistic with the overall skin 
colouration being a light yellowish. All fins in the 
preserved specimen have a light brownish appear- 
ance, apparently internal and possibly an artefact of 
preservation. The dorsal fin spines are blackish at 
their bases. The eyes are black. There is a rectangu- 
lar patch on the dorsal head surface originating just 
posterior to the level of the spiracles and extending 
back over the pectoral fin. It becomes reticulated at 
the rear, and measures 71.6 mm in length and 
24.5 mm in maximum width. An oval spot 5.9 mm 
long is at the first dorsal fin origin and a similar spot 
5.6 mm long at the second dorsal fin origin. There is 



120 



2001 



Notes 



121 




Figure 1. Dorso-lateral view of a rare leucistic Spiny Dogfish, Squalus acanthias, captured about 2 km off Boars Head, 
Bay of Fundy, Nova Scotia, Canada at 44°23'N, 66°15'W, by Stewart Taylor, Captain of the longhner Nadia C, of 
Centreville, Digby County, on 22 May 1989. 



an elongate blotch 9.7 mm long on the left caudal 
peduncle upper margin somewhat behind the caudal 
fin origin, matched by a similar blotch 10.0 mm long 
on the right side but somewhat posterior to the one 
on the left; and a blotch, about 15 mm wide, con- 
cealed in each pectoral fin axil. 

Fr0iland (1975) provides the only other confirmed 
record of a leucistic Spiny Dogfish, taken in the 
North Sea in the late 1960s (with a verbal record of 
an earlier capture). This specimen was a female, 990 
mm total length. Captain Taylor and other fishermen 
in the area of capture of the Nova Scotian specimen 
confirm that this is the first "pure white" dogfish 
seen. Normal-coloured dogfish were numerous in the 
area at the time of capture of the leucistic specimen. 

Acknowledgments 

We are indebted to Captain Taylor who captured 
this Spiny Dogfish, and to Claire Doucette, Fisheries 
Officer, Dibgy for bringing it to the Nova Scotia 
Museum. The photograph was taken by Ron 
Merrick, Media Services, Nova Scotia Museum. 

Literature Cited 

Ben Brahim, R. 1998. Albinisme chez une torpille ocel- 
lee. Torpedo (Torpedo) torpedo. Cybium 22: 83-86. 



Compagno, L. J. V. B. 1984. FAO Species Catalogue. 
Sharks of the World. An Annotated and illustrated cata- 
logue of shark species known to date. Food and 
Agriculture Organization, Rome, Fisheries Synopsis, 
125, Volume 4: viii + 249 pages. 

Dawson, C. E. 1964. A bibliography of anomalies of fish- 
es. Gulf Research Reports 1: 308-399. 

Dawson, C. E. 1966. A bibliography of anomalies of fish- 
es. Supplement 1. Gulf Research Reports 2: 169-176. 

Dawson, C. E. 1971. A bibliography of anomalies of fish- 
es. Supplement 2. Gulf Research Reports 3: 215-239. 

Dawson, C. E., and E. Heal. 1976. A bibliography of 
anomalies of fishes. Supplement 3. Gulf Research 
Reports 5: 35^1. 

Fr0iland, 0. 1975. Albinisme hos hai [Albinism in sharks]. 
Fauna, Oslo 28: 170-173. 

Gopalan, U. K. 1971. On two abnormal sharks from 
Gujarat. Journal of the Bombay Natural History Society 
68: 465^67. 

Lincoln, R. J., G. A. Boxshall, and P. F. Scott. 1982. A 
dictionary of ecology, evolution and systematics. 
Cambridge University Press, Cambridge, viii + 298 pages. 

Talent, L. G. 1973. Albinism in embryo gray smooth- 
hound sharks, Musteliis californicus, from Elkhorn 
Slough, Monterey Bay, California. Copeia 1973: 595- 
597. 

Received 15 May 2000 
Accepted 1 1 February 2002 



122 



The Canadian Field-Naturalist 



Vol. 116 



Anomalies in Incisor Wear of American Elk, Cervus elaphus, 
in the French River Delta, Ontario 



J. Hamr', F. F. Mallory2, I. A. Filioni, G. S. Brown2 and M. A. Jost2 

'Northern Environmental Heritage Institute, Cambrian College of Applied Arts and Technology, 1400 Barrydowne Road, 

Sudbury, Ontario P3A 3V8 Canada 
^Department of Biology, Laurentian University, Sudbury, Ontario P3E 2C6 Canada (Correspondence should be addressed 

to fmallory@nickel.laurentian.ca) 

Hamr, J., F. F. Mallory, I. A. Filion, G. S. Brown, and M. A. Jost. 2002. Anomalies in incisor wear of American Elk, 
Cervus elaphus, in the French River delta, Ontario. Canadian Field-Naturalist 1 16(1): 122-123. 

This article reports on limited observations of dental characteristics of American Elk introduced to the Burwash/French 
River area of Ontario from Alberta in the 1930s. Incisor wear was examined in 29 elk during a four-year study of two, geo- 
graphically distinct herds. All examined adult elk from the French River delta (six females and four males) had excessive 
front incisor (I-l, 1-2) wear as compared to adult animals from the Burwash region (five females, one male). No wear was 
ascertained in calves and yearlings belonging to the French River herd. Radio-monitoring of winter foraging behaviour of 
elk in the French River delta revealed frequent use of rock tripe (Umbilicaria spp.), scraped from cliff faces along water- 
way shorelines. 

Key Words: American Elk, Cervus elaphus, incisor wear anomaly, Ontario. 



Historically, the eastern American Elk (Wapiti) 
inhabited most of southern Ontario and south-western 
Quebec. However, due to over-hunting and habitat 
degradation, elk were extirpated from the eastern half 
of the continent by the late 1800s (Bosveld 1996*). 
The small elk population residing between the north 
shore of Georgian Bay, Lake Huron and Sudbury, 
Ontario (46.00°N, 80.50°W) are remnant animals 
from a 1930s introduction of Rocky Mountain elk 
from Alberta (Polziehn et al. 1998). For at least 30 
years, these Ontario elk have existed in two relatively 
distinct herds, one in the French River delta on 
Georgian Bay (50 km south of Sudbury), and the 
other in the vicinity of the former Burwash Industrial 
Farm, 20 km south of Sudbury (Ranta 1979; Brown 
1998). This area is located within the Great Lakes-St. 
Lawrence ecotone and associated with mixed conifer 
and hardwood forests (Rowe 1972). The climate in 
this region is continental with some moderating influ- 
ences along Georgian Bay. 

As a part of a wider investigation of the remnant 
elk herds, 16 animals were captured and radio-col- 
lared between 1994 and 1997 to determine habitat 
use and causes of mortality. Field necropsies were 
performed on all deceased animals (collared and 
non-collared) that could be located during the study. 
In total, the incisors of 29 captured and dead animals 
were examined (Table I ). 

It was readily apparent that the incisor wear pat- 
tern differed between the two herds (Figures I and 
2). The French River animals had severely worn 
lower incisors, especially I-l and 1-2. Animals 
between 6 and 15 years of age showed the highest 
amount of wear; however, no abrasion of lower 
incisors was apparent in calves nor yearlings. Elk 
replace their deciduous I-l at 15 months of age and 



1-2 at 18 months (Bubenik 1982). Monitoring of 
winter foraging behaviour of instrumented animals 
revealed frequent scraping of lichens, particularly 
rock tripe, from steep cliff faces along the French 
River delta waterways. The use of lichens coincided 
with increased browsing on conifers during late win- 
ter (February-March). Among the consumed conifers 
were Common Juniper (Juniperus communis). 
Eastern White Cedar (Thuja occidentalis), and White 
Pine (Pinus strobus) (Jost et al. 1999). Animals from 
the Burwash herd also consumed conifers, but 
showed normal incisor wear. The variation in incisor 
wear appeared to result from differences in regional 
topography. The Burwash region had more rolling 
terrain, fewer cliff faces, and fewer waterways as 
compared to the French River delta. 

It is generally known that lichens comprise a large 
portion of the winter diet of Caribou and Reindeer 
(Rangifer tarandus) and are high in digestible carbo- 
hydrates and low in nitrogen (Thomas et al. 1984). 
Elk wintering in mature conifer forests in the Rocky 
Mountains readily consumed arboreal lichens (Hash 
1973; Bohne 1974). Usnic acid is a common sec- 
ondary compound found in lichens (Culberson 1977) 
and data collected by Palo (1993) suggested that the 
addition of usnic acid to hay resulted in higher in 



Table 1. Nii 
and 2 herds. 


mber of examined 


animals 


in 3 age classes 


Herd 


calves 
(0.5-1 year) 


yearlings 

(1.5-2 years) 


adults 

(2.5-15 years) 


Burwash 
French River 


6 

1 




2 
4 


6 
10 



2002 



Notes 



123 




Figure 1 . Lower incisors of adult elk from the French River 
population, near Sudbury, Ontario showing severe 
wear associated with scraping lichens from cliff 
faces. 



vitro dry matter digestibility. Jenks and Leslie (1988) 
also found that in vitro digestion of White-tailed 
Deer {Odocoileus virginianus) forage was enhanced 
by lichens, increasing fermentation efficiency and 
forage digestibility. In the present study, use of 
lichens coincided with high snow depths and 
increased utilization of conifers during late winter 
(February and early March) and to a lesser extent 
early spring (late March and April). It is suggested 
that where available, rock tripe consumption 
enhanced the digestibility of conifers and other 
woody browse consumed by the studied elk in late 
winter and contributed to the observed variation in 
tooth wear patterns. 

Acknowledgments 

We thank the Rocky Mountain Elk Foundation, 
Safari Club International (Ontario Chapter). Tembec 
Forest Products. Ontario Federation of Anglers and 
Hunters, Ontario Ministry of Natural Resources, 
Department of National Defense, Parks Canada, 
Sudbury Fish and Game Protective Association. 
Cambrian College, and Laurentian University for pro- 
viding financial support. Heli North Aviation, Atwood 
Island Lodge, and Hartley Bay Marina. J. F. 
Zuchlinski, and T. L. Hillis, provided technical sup- 
port. 

Literature Cited 

Bosveld, H. J. 1996. A review of documented occur- 
rences of native elk (Cervus elaphus) in Ontario. 
Heritage Consultants. Parks Canada. Ontario Region. 
Cornwall, Ontario. 26 pages. 



Figure 2. Lower incisors of adult elk from the Burwash 
population, showing a normal wear pattern. 



Brown, G. S. 1998. Spatial behaviour and habitat utiliza- 
tion by wapiti (Cervus elaphus) in the French River and 
Burwash regions of Ontario. M.Sc. thesis. Laurentian 
University, Sudbury. 77 pages. 

Bubenik, A. B. 1982. Physiology. Pages 125-179 /« Elk 
of North America — Ecology and Management. Edited 
by J. W. Thomas and E. Toweill. Wildlife Management 
Institute. Harrisburg. 698 pages. 

Culberson, C. 1977. Chemical and botanical quick guide 
to lichen products. Bryologist 73: 77-377. 

Hash, H. S. 1973. Movements and food habits of the 
Lochsa elk. M.S. thesis, University of Idaho. Moscow. 
85 pages. 

Jenks, J. A., and D. M. Leslie. 1988. Effect of lichen and 
in vitro methodology on digestibility of winter deer diets 
in Maine. Canadian Field-Naturalist 102: 216-220. 

Jost, M. A., J. Hamr, L Fillon, and F. F. Mallory. 1999. 
Forage selection by elk in habitats common to the 
French River/Burwash region of Ontario. Canadian 
Journal of Zoology 77: 1429-1438. 

Palo, R. T. 1993. Usnic acid, a secondary metabolite of 
lichens and its effect on in vitro digestibility in reindeer. 
Rangifer 13: 39-43. 

Polziehn, R. C, J. Hamr, F. F. Mallory, and C. Strobeck. 
1998. Phylogenetic status of North American wapiti 
(Cervus elaphus) subspecies. Canadian Journal of 
Zoology 76: 998-1010. 

Ranta, B. 1979. Range and habitat relationships of wapiti 
(Cervus canadensis) in the Burwash-French River area 
of Ontario. M.Sc. thesis. Carleton University. Ottawa. 
205 pages. 

Rowe, J. S. 1972. Forest regions of Canada. Publication 
number 1300. Canadian Forest Service. Ottawa. 

Thomas, D. C, P. Kroeger, and D. Herviux. 1984 //; 
vitro digestibilities of plants utilized by barren ground 
caribou. Arctic 37: 31-36. 

Received 24 March 2000 
Accepted 4 April 2tK)2 



124 



The Canadian Field-Naturalist 



Vol. 116 



First record of the Hoary Bat, Lasiurus cinereus (Chiroptera: 
Vespertilionidae), from Prince Edward Island. 

Donald F. McAlpinei, Frances Muldoon^, and Alexander I. Wandeler^ 

'Natural Science Department, New Brunswick Museum, 277 Douglas Avenue, Saint John, New Brunswick E2K 1E5 

Canada; e-mail: dmcalpin@nb.aibn.com 
2Canadian Food Inspection Agency, Animal Diseases Research Institute, 3851 Fallowfield Road, P.O. Box 11300, Station 

H, Nepean Ontario K2H 8P9 Canada. 

McAlpine, Donald P., Frances Muldoon, and Alexander I. Wandeler. 2002. First record of the Hoary Bat, Lasiurus 
cinereus (Chiroptera: Vespertilionidae), from Prince Edward Island. Canadian Field-Naturalist 1 16(1): 124-125. 

The Hoary Bat, Lasiurus cinereus, is reported from Prince Edward Island for the first time on the basis of an adult animal 
collected 17 August 1999 and submitted for rabies testing. Based on previous records for this species from the Atlantic 
region, the presence of the Hoary Bat on Prince Edward Island is not unexpected. Although Lasiurus cinereus appears to 
be rare in the province, bat surveys that nndght better establish the species status on the Island are lacking. 

Key Words: Hoary Bat, Lasiurus cinereus, new record. Prince Edward Island. 



The Hoary Bat, Lasiurus cinereus, is the most 
widespread of North American bats, and although 
infrequently encountered, is widely distributed in 
Canada (van Zyll de Jong 1985). Maunder (1988) 
summaiized the 17 records for Atlantic Canada up to 
that time, including the first occurrence from New- 
foundland. Previously, Lasiurus cinereus has not 
been recorded on Prince Edward Island, and has 
been considered hypothetical for Kouchibouguac 
National Park, which borders the coast of New 
Brunswick adjacent to the Island (Maunder 1988; 
Tremblay 1992). However, recent acoustic surveys 
revealed the species presence in the park (H. 
Broders, personal communication to DFM). Based 
on these earlier reports from the region, the presence 
of Lasiurus cinereus on Prince Edward Island was 
not unexpected. 

The first specimen of the Hoary Bat on Prince 
Edward Island was collected 17 August 1999, at 
Charlottetown, Queens County, and submitted to the 
Canadian Food Inspection Agency for rabies testing. 
Although the rabies specimen submission form 
records that the animal was aggressive and was 
killed, it was negative for rabies virus. Only the head 
was submitted and it has now been deposited in the 
New Brunswick Museum collection (NBM 5801). 
The head includes the skull, with cranium slit, of an 
adult animal with pelage intact across the muzzle 
and the lower jaws posterior to the eyes. 

Nearly all records for the Hoary Bat in Atlantic 
Canada have been individuals encountered in late 
summer and autumn, with the Prince Edward Island 
record reported here following this pattern. Maunder 
(1988) suggested that late autumn tree bats may be 
present in the Atlantic region as a result of "drift 
migration" and van Zyll de Jong (1985) noted that 
this species has been recorded from widely scattered 
locations far beyond areas that appear to offer suit- 
able habitat. Although Lasiurus cinereus appears to 



be rare on Prince Edward Island, bat surveys that 
might better establish the species status on the Island 
are lacking. 

To date there is no evidence that the Hoary Bat 
breeds in the Atlantic region. Although there are two 
mid- July occurrences for Quebec (Pierre Aquin, per- 
sonal communication to DFM) there are only two July 
records for Maritime Canada, both for New 
Brunswick (12 and 29 July, NBM 1163, Tremblay 
1989*). All other reports of the Hoary Bat from the 
Atlantic Provinces are from early August to 17 
November. 

Acknowledgments 

For sharing current information on the status of 
the Hoary Bat in the Atlantic region with us we 
thank Pierre Aquin, Quebec Natural Heritage Data 
Centre; Kate Bredin, Atlantic Canada Conservation 
Data Centre; Hugh Broders, New Brunswick 
Cooperative Fish and Wildlife Research Unit, Uni- 
versity of New Brunswick; and Andrew Hebda, 
Nova Scotia Museum of Natural History. 

Documents Cited [marked* in text] 

Tremblay, E. 1989. A brief survey of the chiropterian 
fauna of Fundy National Park. Unpublished report. Parks 
Canada, Natural Resources Conservation, Fundy 
National Park, 19 pages plus appendices. 

Literature Cited 

Maunder, J. E. 1988. First Newfoundland record of the 
Hoary Bat, Lasiurus cinereus, with a discussion of other 
records of migratory tree bats in Atlantic Canada. 
Canadian Field-Naturalist 102: 726-728. 

Tremblay, E. 1992. Bats of Kouchibouguac and Fundy 
National Parks, New Brunswick, Canada. Pages 
291-294 in Science and the Management of Protected 
Areas. Edited by J. Willison, S. Bondrup-Nielson, C. 
Drysdale, T. B. Herman, N. W. P. Munro, and T. L. 
Pollock. Elsevier, Amsterdam. 



2002 



Notes 



125 



van Zyll de Jong, C. G. 1985. Handbook of Canadian 
Mammals Number 2: Bats. National Museum of Natural 
Sciences, Ottawa, 212 pages. 



Received 30 May 2000 
Accepted 24 January 2002 



Wolf, Canis lupus, Response to Domestic Sled Dog, Canis familiaris. 
Activities in Central Yukon 



Gerald W. Kuzyk^'^ and Kristin M. Kuzyk^ 



'Department of Renewable Resources, University of Alberta, Edmonton, Alberta T6G 2H1 Canada 

^Faculty of Science, University of Alberta, Edmonton, Alberta T6G 2H1 Canada 

^School of Environmental Sciences, Lakeland College, Vermilion, Alberta T9X 1K5 Canada 

Kuzyk, Gerald W., and Kristin M. Kuzyk. 2002. Wolf, Canis lupus, response to domestic sled dog, Canis familiaris, activi- 
ties in central Yukon. Canadian Field-Naturalist 116(1): 125-126. 

We present an observation of a lone Wolf {Canis lupus) regularly following teams of domestic sled dogs {Canis familiaris) 
and humans for distances up to 34 kilometers. The lone Wolf died of unknown causes, approximately 30 meters from a 
trapping trail used by sled dog teams and within vocalization distance of sled dogs at an occupied cabin. We also noted a 
pack of about seven Wolves that did not follow the same sled dog trail when they encountered it, and exchanged vocaliza- 
tions with the dogs. We suggest human-made trails can be especially important travel corridors for lone Wolves. 

Key Words: Wolf, Canis lupus. Dog, Canis familiaris. Black Bear, Ursus americanus. Lake Whitefish, Coregonus 
clupeaformis, travel corridor, Yukon, howling, communication. 



Wolves {Canis lupus) are ancestors of domestic 
dogs {Canis familiaris) (Morey 1992); are known to 
interbreed with dogs (Maagaard and Graugaard 1994); 
to aggressively interact with, and prey upon dogs 
(Tompa 1983); and to closely follow travelling sled 
dog teams (Karras 1975). Coppinger and Coppinger 
(1995) reviewed literature of livestock guard dogs in 
Europe and Asia, and applied this information to a 
study of Wolves and guard dogs in Minnesota, con- 
cluding that Wolves can treat dogs as conspecifics and 
not as prey. Reports of Wolf-dog interactions in the 
scientific literature are scarce (Fritts and Paul 1989), 
especially in areas of low human activity. In Yukon, 
most known Wolf-dog encounters are predatory, and 
usually occur in autumn and winter (Alan Baer, 
Yukon Fish and Wildlife Branch, personal communi- 
cation). 

From September 1991 to February 1992 we used 
13 sled dogs for the purpose of trapping, south of 
Mayo in central Yukon (63° 30' N 135° 30' W). On 9 
October 1991, at approximately 09:00 and 12:00 h, a 
pack of Wolves was heard howling within a couple 
kilometers of our trapline cabin. There was no 
response from the sled dogs that were chained sepa- 
rately within 40 meters of the trapline cabin. On 10 
October a small Wolf track, presumably that of a 
pup, was noticed in the fresh snow within ten meters 
of a sled dog chained at the outside edge of the 
group of dogs. Two of the dogs had barked that 
night, but the cause may have been a Black Bear 
{Ursus americanus), as its tracks in fresh snow were 



noticed within 75 meters of the cabin. On 12 
October, about two kilometers from the cabin, we 
discovered a Lake Whitefish {Coregonus clupea- 
formis) spawning run in a small narrows of less than 
five meters wide that was being used by a pack of 
Wolves and a Black Bear. Wolf and Black Bear 
tracks were obvious in the fresh snow near the creek. 
Whole and partially eaten Whitefish, as well as a 
cache of buried Whitefish, were found on the creek 
bank. The Wolves remained near the fish run for 
seven days until 19 October when the ice thickened 
and the fish run neared completion. 

On 8 December at 10:45 h, excited barking from 
the sled dogs drew our attention to a lone gray Wolf 
on the lake ice approximately 400 meters from our 
cabin. It quickly fled into the forest when it noticed 
humans near the cabin. The same afternoon, a single 
fresh Wolf track was seen on the main trapping trail, 
which had initially been made with snowshoes, and 
was regularly used for travel with sled dogs. A lone 
Wolf traveled on the trapping trail for approximately 
five kilometers and showed interest in a trap set for 
Wolverine {Gulo gulo) that had a Moose {Alces 
alces) head for bait. On the morning of 1 1 December 
a single large fresh Wolf track was noticed within 30 
meters of a sled dog chained at the periphery oi the 
group of dogs. A Wolf remained in the area of the 
cabin, successfully taking bait from traps and scav- 
enging cached Whitefish at the creek narrows for the 
next couple days. On 14 December a Wolf was 
caught in and then escaped from a #4 long spring 



126 



The Canadian Field-Naturalist 



Vol. 116 



trap, set for Lynx {Lynx canadensis). The same 
evening from approximately 19:00-19:15 h, howling 
from a single Wolf was heard across the lake from 
the cabin, with no reply from the dogs. 

In late December and early January, lone Wolf 
tracks were a frequent occurrence on and near the 
main trapping trail. On the morning of 17 January 
1992, we left the main cabin and traveled 17 kilome- 
ters to a smaller line cabin. During the night there 
was a light snowfall and the following morning, we 
noticed a single fresh Wolf track 30 meters from the 
cabin. A female sled dog in estrous was tied near the 
front door of the cabin, and no barking by the dogs 
was heard at night when the Wolf was nearby. On 19 
January, during the return trip to the main cabin, we 
encountered a single fresh Wolf track in the snow, 
which suggested the same lone Wolf had followed us 
to the small line cabin. On 20 January, a fresh lone 
Wolf track was observed within two kilometers of 
the main cabin, suggesting the same individual Wolf 
had followed us on the return trip back to the main 
cabin, a distance totaling approximately 34 kilome- 
ters. Wolves maintain their tenitories by killing tres- 
passing Wolves (Mech 1991), so during this foray, 
the lone Wolf had traveled well inside an occupied 
Wolf pack territory and risked being killed by terri- 
torial Wolves. 

On 23 January at about 10:00 h, and approximate- 
ly three kilometers from the main cabin, excitement 
from the team of sled dogs alerted us to a dead Wolf 
in its bed about 30 meters from the trapping trail. We 
assume it was the same Wolf that had been in the 
locality, due to the gray pelage and the fact that fol- 
lowing this discovery, we saw no more evidence of 
lone Wolf tracks on the trails and heard no more 
howling from a single Wolf. The cause of the Wolfs 
death was undetermined; there were no signs of 
external injuries from other Wolves and rigor mortis 
had not occurred. The Wolf was an old male as 
determined by tooth wear, and was in an emaciated 
state, as indicated by the lack of body fat and pro- 
truding ribs, although the coat was still in prime con- 
dition. We believe the site of death for this Wolf was 
not random because it was only a short distance from 
the compacted trail the Wolf had used in recent trav- 
els. The site was also within vocalization distance of 
sled dogs and in view (three kilometers across a val- 
ley) of the occupied cabin. Human-made trails can 
provide easier movement for Wolves (James 1999) 
and as in this case, may be especially important for 
lone Wolves in poor condition. 

On 25 January 1992 a pack of an estimated seven 
Wolves encountered our trapping trail on a narrow 
creek, the same one used by the lone Wolf. There 
was little snow at this location and the Wolves 
fanned out and did not follow the trail off the creek. 
On 2 February at approximately 12:00 h, while we 
were resting the two dog teams near the same loca- 



tion, a pack of Wolves began howling approximately 
five kilometers away. The air was still, the sky sunny 
and the temperature estimated at -20C. All the dogs 
became attentive, looked in the direction of the 
Wolves and began synchronously howling for 
approximately one minute. They then became silent 
and attentive, looking in the Wolves' direction. The 
Wolves and sled dogs exchanged howls once more, 
then there was prolonged silence and we left the 
area. Wolves howl as a means of territorial mainte- 
nance and as communication within and between 
packs (Harrington and Mech 1979). No scientific 
reference was found of Wolves and dogs communi- 
cating through howling, so this observation may help 
in further understanding Wolf-dog interactions. 

Acknowledgments 

We thank Bernard Menelon and Barry Pouche 
both of Mayo, Yukon, for allowing us the use of 
their trapline. Ludwig Carbyn (Canadian Wildlife 
Service, Edmonton, Alberta) and Alan Baer (Fish 
and Wildlife Branch, Whitehorse, Yukon) kindly 
reviewed this note and added helpful suggestions. 

Literature Cited 

Coppinger R., and L. Coppinger. 1995. Interactions 
between livestock guarding dogs and wolves. Pages 
523-526. In Ecology and conservation of wolves in a 
changing world. Edited by L. N. Carbyn, S. H. Fritts and 
D. R. Seip. Canadian Circumpolar Institute, University 
of Alberta, Edmonton. 

Fritts, S. H., and W. J. Paul. 1989. Interactions of wolves 
and dogs in Minnesota. Wildlife Society Bulletin 17: 
121-123. 

Harrington, F. H., and L. D. Mech. 1979. Wolf howling 
and its role in territory maintenance. Behaviour 68: 
207-249. 

James, A. R. C. 1999. Effects of industrial development 
on the predator-prey relationship between wolves and 
caribou in northeastern Alberta. Ph.D. thesis. University 
of Alberta, Edmonton, Alberta. 70 pages. 

Karras, A. L. 1975. Face the north wind. Burns and 
Maceachem Limited, Don Mills, Ontario. 191 pages. 

Maagaard, L., and J. Graugaard. 1994. Female arctic 
wolf, Canis lupus arctos, mating with domestic dogs, 
Canis familiaris, in northeast Greenland. Canadian 
Field-Naturalist 108: 374-375. 

Mech, L. D. 1991. The way of the wolf Voyageur Press, 
Stillwater, Minnesota, 120 pages. 

Morey, D. F. 1992. Size, shape and development in the 
evolution of the domestic dog. Journal of Archeological 
Science 19: 181-204. 

Tompa. 1983. Problem wolf management in British 
Columbia: conflict and program evaluation. Pages 112- 
1 19 in Wolves in Canada and Alaska: their status, biolo- 
gy, and management. Edited by L. N. Carbyn in 
Canadian Wildlife Service Report Series, Number 45. 

Received 10 June 2000 
Accepted 22 February 2002 



I 



2002 



Notes 



127 



First Record of an Eastern Coyote, Canis latrans, in Labrador 

Tony E. Chubbs^ and Frank R. Phillips^ 

^Department of National Defence, 5 Wing Goose Bay, Box 7002, Station A, Happy Valley-Goose Bay, Labrador, 

Newfoundland AOP ISO, Canada 
^Department of Forest Resources and Agrifoods, Government of Newfoundland and Labrador, Box 3014, Station B, Happy 

Valley-Goose Bay, Labrador, Newfoundland AOP lEO Canada 

Chubbs, Tony E., and Frank R. Phillips. 2002. First record of an eastern Coyote, Canis latrans, in Labrador. Canadian 
Field-Naturalist 116(1): 127-129. 

An adult male Eastern Coyote, Canis latrans, trapped on 14 January 1995 along the Churchill River, is the first confirmed 
record for Labrador. This record is approximately 600 km north east of the previously accepted range limit in eastern 
Canada. 

Key Words: Eastern Coyote, Canis latrans, range, distribution, Labrador. 



Coyotes {Canis latrans) have been expanding 
their range northward and eastward in North 
America during the past century (Perkins and Mautz 
1990; Thurber and Peterson 1991; Lehman et al. 
1991; Parker 1995). Coyotes were first recorded in 
Quebec in 1944 (Rand 1945) and had spread north 
across the Gaspe Peninsula by 1974 (Georges 1976). 
More recently, Coyotes have become established as 
a significant component of the boreal forests of 
southeastern Quebec (Crete and Desrosiers 1995; 
Tremblay et al. 1998; Patterson et al. 1999). Coyotes 
immigrated to the island of Newfoundland on drift- 
ing pack ice and were first documented in 1985 
(Parker 1995). Here, we report the first record of an 
Eastern Coyote in Labrador, approximately 600 km 
north east of the previously recorded range limit in 
Quebec (George 1976; Parker 1995). 

On 14 January 1995, an adult male Coyote was 
trapped along the Churchill River (53°17'N, 
60° 15' W), approximately 2 km south east of Happy 
Valley-Goose Bay. The Coyote was estimated (F. 
Blake, personal communication) to weigh approxi- 
mately 15 kg. Although the weight could not be con- 
firmed, it is consistent with reported average weights 
of 13.5 kg, and 12.5 kg for adult male forest Coyotes 
in Quebec (Tremblay et al. 1998; Poulle, M.-L. et al. 
1995) and about half the weight (ca. 30 kg) of 
Wolves {Canis lupus) in Labrador (Parker and 
Luttich 1986). Dr. C. G. van Zyll de Jong of the 
National Museum of Natural Sciences, later exam- 
ined the skull and performed a quantitative compari- 
son of the Goose Bay skull with samples of Coyotes 
and Wolves of both sexes using a ratio diagram 
(Figure 1). Eleven different dimensions of the skull 
(see Nowak 1979) for each of the taxa and the Goose 
Bay specimen were compared. All measurements 
were converted into logarithms and the sample of 
male Wolves was used as the standard. The differ- 
ence (d) between the logarithmic value of the stan- 
dard and the other groups and the Goose Bay speci- 
men were calculated, plotted and connected by lines 



to aid in their interpretation (Figure 1). This compar- 
ison revealed that the Goose Bay skull ratio series 
was similar to those of the male and female Coyote 
samples and within two standard deviations from the 
male Coyote sample means. These results confirmed 
its identification as that of a large adult male Coyote 
(Dr. C. G. van Zyll de Jong, 17 December 1995, per- 
sonal communication). Recent information on 
expanding Coyote populations in eastern Quebec 
also indicates that Coyotes appear to be larger in the 
eastern portion of their range (Thurber and Peterson 
1991; Lariviere and Crete 1993). 

Although no previous records of Coyotes exist for 
Labrador, the possibility of an established population 
remains plausible, as they do coexist with Wolves in 
Alaska (Thurber and Peterson 1991), Alberta, British 
Columbia, and more recently Idaho (Pilgrim et al. 
1998). No geographic barriers seem to exist to pre- 
vent Coyotes from extending their range northward 
into Labrador. Delineation of Coyote range expan- 
sion in Labrador may have been precluded by the 
fact that no systematic surveys have been conducted 
for any carnivore species in Labrador and low fur 
prices in the last decade have significantly decreased 
trapping effort resulting in only one specimen to 
date. The expansion of Coyotes into eastern forests 
has occurred due to high populations of White-tailed 
Deer {Odocoileus virginianus), high populations of 
humans and associated garbage, and changes in for- 
est communities, all of which have resulted in an 
improvement in food supply (Thurber and Peterson 
1991; Tremblay et al. 1998). Both sedentary and 
migratory Caribou {Rangifer tarandus) herds fre- 
quent central and southern Labrador and Coyotes 
have been identified as a threat to remnant popula- 
tions of Woodland Caribou in southeastern Quebec 
(Crete and Desrosiers 1995). Recent extensions of 
the winter range by migratory Caribou from the 
George River Caribou herd (Department of National 
Defence, unpublished data), and increased numbers 
of Moose {Alces alecs) in southern Labrador 



128 



The Canadian Field-Naturalist 



Vol. 116 



• C. Latrans (males, n=166) 
M C. latrans (females, n=111) 
A — Goose Bay Specimen 
— O— -C. lupus (females, n=146) 
— Q — C. lupus (males, n=233) 
- - #■ - Standard deviation C. lupus (male) 
------ Standard deviation C. latrans (male) 



-0.2 



II CO 




-0.1 A' 



1.GSL 



2. ZW 



6. PW 



10. LTB 



11. HMO 



12. DJ 



13. DC 



14. LP4 



15. WM1 



Skull 
Dimension 

Figure 1 . Ratio diagram comparing eleven standard cranial measurements of the Goose Bay, Labrador specimen with sam- 
ples of male and female Canis lupus and C. latrans. For a description of measurements and samples of C. lupus 
and C. latrans see: Appendix B in Nowak (1979). 



(Chubbs and Schaefer 1997) may provide a suffi- 
cient prey and carrion base to support Coyotes 
through the winter. Coyotes are a highly adaptable 
species (Pilgrim et al. 1998) and are capable of 
remaining in an area with a seasonally reduced prey 
base and harsh weather conditions (Patterson et al. 
1999). Although the boreal forest appears to be a rel- 
atively poor habitat for Coyotes (Tremblay et al. 
1998), Coyotes can persist at low densities, depend- 
ing mainly on Snowshoe Hares (Lepus americanus), 
mice and voles (Patterson et al. 2000; M. Crete per- 
sonal communication). It has been known for some 
time (Lehman et al. 1991; Pilgrim et al. 1998) that 
hybridisation of Wolf and Coyote genotypes has 
already occurred in Quebec due to the rapid north- 
east progression of Coyotes. 

Although this single record may not confirm a 
range extension, the potential for Coyote range 
expansion into Labrador appears to be plausible and 
even likely. The sparse human population of 
Labrador, lack of systematic surveys, and decreas- 
ing trapping effort may result in an existing sparse 



population going undetected. Despite the presence 
of a healthy Wolf population, which may limit or 
slow down colonisation through intraguild preda- 
tion, the increasingly human-transformed landscape 
may be the key to the establishment of Coyotes in 
Labrador. 

Acknowledgments 

We appreciate the support of the Newfoundland- 
Labrador Wildlife Division, Goose Bay, Labrador 
and J. A. Schaefer, who arranged for the skull mea- 
surements and identification conducted by the late 
C. G. van Zyll de Jong. Thanks are extended to trap- 
per Fred Blake for providing the skull and details of 
the capture location and to D. Laing for drafting the 
figure. Funding for this publication was provided 
through the Environmental Monitoring Program, 
through the Department of National Defence (DND). 
We thank DND (Major G. Humphries) for their con- 
tinued support of wildlife research in Labrador. We 
also thank M. Crete and two anonymous referees for 
commenting on an earlier draft of this manuscript. 



2002 



Notes 



129 



Literature Cited 

Chubbs, T. E., and J. A. Schaefer. 1997. Population 
growth of Moose, Alces alces, in Labrador. Canadian 
Field-Naturalist 111: 238-242. 

Crete, M., and A. Desrosiers. 1995. Range expansion of 
Coyotes, Catiis latrans, threatens a remnant herd of 
Caribou, Rangifer tarandus, in southeastern Quebec. 
Canadian Field-Naturalist 109: 227-235. 

Georges, S. 1976. A range extension of the Coyote in 
Quebec. Canadian Field-Naturahst 90: 78-79. 

Lariviere, S., and M. Crete. 1993. The size of eastern 
Coyotes {Canis latrans): a comment. Journal of Mam- 
malogy 74: 1072-1074. 

Lehman, N., A. Eisenhawer, K. Hansen, L. D. Mech, 
R. O. Peterson, P. J. P. Gogan, and R. K. Wayne. 
1991. Introgression of coyote mitochondrial DNA into 
sympatric North American Grey Wolf populations. 
Evolution 45: 104-119. 

Nowak, R. M. 1979. North American Quaternary Canis. 
Monograph of the Museum of Natural History, Univer- 
sity Arkansas, number 6, Arkansas. 

Parker, G. R. 1995. Eastern Coyote: The story of its 
success. Nimbus Publishing Ltd., Halifax, Nova Scotia. 
256 pages. 

Parker, G. R., and S. Luttich. 1986. Characteristics of 
the Wolf {Canis lupus labradorius Goldman) in 
Northern Quebec and Labrador. Arctic 39: 145-149. 

Patterson, B. R., S. B.-Nielsen, and F. Messier. 1999. 
Activity patterns and daily movements of the Eastern 



Coyote, Canis latrans, in Nova Scotia. Canadian Field- 
Namralist 113: 251-257. 

Patterson, B. R., L. K. Benjamin, and F. Messier. 2000. 
Winter nutritional condition of eastern coyotes in rela- 
tion to prey density. Canadian Journal of Zoology 78: 
420-427. 

Perkins, P. J., and W. W. Mautz. 1990. Energy require- 
ments of eastern coyotes. Canadian Journal of Zoology 
68: 656-659. 

Pilgrim, K. L., D. K. Boyd, and S. H. Forbes. 1998. 
Testing for wolf-coyote hybridisation in the Rocky 
Mountains using mitochondrial DNA. Journal of Wild- 
life Management 62: 683-689. 

Poulle, M.-L., M. Crete, and J. Huot. 1995. Seasonal 
variation in body mass and composition of eastern coy- 
otes. Canadian Journal of Zoology 73: 1625-1633. 

Rand, A. L. 1945. Mammals of the Ottawa district. 
Canadian Field-Naturalist 59: 1 1 1-132. 

Thurber, J. M., and R. O. Peterson. 1991. Changes in 
body size associated with range expansion in the Coyote 
{Canis latrans). Journal of Mammalogy 72: 750-755. 

Tremblay, J. P., M. Crete, and J. Huot. 1998. Summer 
foraging behaviour of eastern coyotes in rural versus for- 
est landscape: A possible mechanism of source-sink 
dynamics. Ecoscience 5: 172-182. 

Received 24 February 2000 
Accepted 25 February 2002 



Long Distance Movement by a Coyote, Canis latrans, and Red Fox, 
Vulpes vulpes, in Ontario: Implications for Disease-spread 

Richard C. Rosatte 

Ontario Ministry of Natural Resources, Wildlife Research and Development Section, Trent University, Science Complex. 
P.O. Box 4840, Peterborough, Ontario, K9J 8N8 Canada 

Rosatte, Richard C. 2002. Long distance movement by a Coyote, Canis latrans, and Red Fox, Vulpes vulpes, in Ontario: 
Implications for disease-spread. Canadian Field-Naturalist 116(1): 129-131. 

During a rabies control program in southern Ontario, Raccoons, {Procyon lotor lotor). Striped Skunks {Mephitis mephitis). 
Red Foxes {Vulpes vulpes), and Coyotes {Canis latrans) were live-captured, vaccinated, ear-tagged, and released at point 
of capture. One of eight Coyotes captured and released during 1995 in Niagara Falls, Ontario, dispersed 320 km to 
Coatsworth, Ontario. Additionally, 1 of 23 foxes, captured and released in Scarborough, Ontario, during 1994, moved 170 
km to Rossmore, Ontario. Although such long distance movements are probably rare in Ontario they may play a critical 
role in the dissemination of infectious diseases such as rabies. 

Key Words: Red Fox, Vulpes vulpes. Coyote, Canis latrans, rabies, movement, dispersal. Ontario. 



As part of a Provincial strategy to prevent raccoon 
rabies from becoming established in Ontario, the 
Ontario Ministry of Natural Resources (OMNR) 
implemented a Trap-Vaccinate-Release (TVR) pro- 
gram in the Niagara Frontier (Rosatte et al. 1997). 
Since 1994, personnel from the OMNR Rabies 
Research Unit have been live-capturing and vacci- 
nating Raccoons {Procyon lotor). Striped Skunks 
{Mephitis mephitis). Red Foxes {Vulpes vulpes) and 



Coyotes {Canis latrans) in a 680 km- area encom- 
passing Niagara Falls, Ontario. During the 1995 
Niagara TVR program, eight Coyotes were live- 
captured, vaccinated, ear-tagged, and released at the 
point of capture in the City of Niagara Falls. One of 
the Coyotes, a juvenile male, released on 26 July 
1995, was shot by a hunter near the town of Coats- 
worth, Ontario, during February 1996; a straight line 
distance of 320 km west (Figure 1). All local 



130 



The Canadian Field-Naturalisx 



Vol. 116 




NEW YORK STATE 



Figure 1. Dispersal direction of a Coyote and a Red Fox in Southern Ontario. 



humane societies and animal control agencies in the 
Niagara TVR area were questioned and none had 
relocated any Coyotes between July 1995 and 
February 1996. 

From 1987-1997, the OMNR Rabies Research 
Unit conducted a TVR program to control rabies in 
Skunks in the city of Scarborough, Ontario. About 
3200 Striped Skunks, 1 1 300 Raccoons and 100 Red 
Foxes were live-captured, vaccinated, ear-tagged, 
and released (Rosatte et al. 1992). During the 1994 
Scarborough TVR program, three Red Foxes were 
captured, vaccinated, ear-tagged, and released at 
point of capture. On 2 January 1996, a trapper cap- 
tured one of the foxes near Rossmore, Ontario; a 
straight line distance of 170 km east (Figure 1). This 
fox, an adult female in 1996, was originally captured 
as a juvenile in Scarborough on 19 August 1994. 

During 1996-1999, 127 Red Foxes and 13 
Coyotes were ear-tagged during TVR operations 
conducted by OMNR in Niagara, Scarborough, and 
Brockville. No significant movements such as those 
noted above were reported. 



Discussion 

In Ontario, juvenile Red Foxes and Coyotes usual- 
ly disperse during their first fall or winter (Voigt 
1987; MacDonald and Voigt 1985; Voigt and Berg 
1987). Researchers in one rural Ontario study deter- 
mined that Coyote movements ranged from 16 km to 
152 km (Kolenosky et al. 1978). Dispersal of juve- 
nile male and female Red Foxes in another study in 
rural Ontario averaged 30 km and 8 km, respective- 
ly. Few individuals dispersed more than 100 km 
(one juvenile male Red Fox dispersed 122 km with 
> 120 total Foxes collared) (Voigt 1987; Voigt and 
MacDonald 1984). Similarly, in Toronto, long dis- 
tance dispersal was rare; only 2 of 33 radio-collared 
Red Foxes dispersed over 100 km (Rosatte unpub- 
lished; Rosatte et al. 1991). 

Although the movements noted in this study 
(320 km for the Coyote and 170 km for the Red Fox) 
are exceptional for Ontario, similar movements have 
been reported elsewhere. For example, Carbyn and 
Paquet (1986) recorded a 544 km movement of an 
adult female Coyote that dispersed from Manitoba to 



2002 



Notes 



131 



Saskatchewan. Additionally, one sub-adult male Red 
Fox moved 395 km from Wisconsin to Indiana 
(Abies 1965). However, such long distance disper- 
sals appear to be rare. 

The last major wildlife rabies epizootic in Canada 
occurred in the Arctic during the 1940s (Tabel et al. 
1974). Numerous wildlife species including Red 
Foxes and Coyotes were reported with rabies as the 
disease dispersed into the Canadian provinces from 
the Northwest Territories in the 1950s (Tabel et al. 
1974). The epizootic reached southern Ontario in 
1956 due mainly to the movements of infected Red 
Foxes (Johnston and Beauregard 1969). Fox rabies 
remained enzootic in southern Ontario since the 
1950s, however, the prevalence of that disease has 
recently declined dramatically due to rabies manage- 
ment programs involving the use of oral rabies vac- 
cine baits and TVR (Maclnnes 1987; Rosatte et al. 
1992; Rosatte et al. 1993). Dispersing Red Foxes and 
to a lesser extent Coyotes (as Red Foxes are the pri- 
mary vector of the Arctic fox strain of rabies) in 
Ontario have likely played a major role in the main- 
tenance of the disease as well as in the initiation of 
new outbreaks (Maclnnes 1987; Johnston and 
Beauregard 1969; Tabel et al. 1974). To minimize 
the risk of dispersing Red Foxes contributing to the 
spread of rabies in Ontario, OMNR is annually dis- 
tributing oral rabies vaccine baits over areas that 
include a 50 km buffer around any rabies case. 
Based on observations from this study, long-distance 
movements by Red Foxes and Coyotes likely are 
rare. However, biologists should consider the poten- 
tial maximum dispersal of Red Foxes and Coyotes 
when planning or implementing a rabies manage- 
ment strategy for Ontario or other areas where either 
of these two species occur. 

Acknowledgments 

Special thanks to Garnie Skinkle and Richard 
Provost for reporting the ear-tagged Red Fox and 
Coyote. The co-operation of the hunters and trap- 
pers of Ontario is critical to the evaluation of the 
OMNR rabies program. C. Davies, K. Abraham, M. 
Obbard, A. Jano, CD. Maclnnes and D. R. Voigt 
reviewed the manuscript. This is OMNR Wildlife 
Research and Development Section contribution 
number 20. 

Literature Cited 

Abies, E. D. 1965. An exceptional fox movement. Journal 
of Mammalogy 46: 102. 



Carbyn, L., and P. Paquet. 1986. Long distance move- 
ment of a coyote from Riding Mountain National Park. 
Journal of Wildlife Management 50: 89. 

Johnston, D., and M. Beauregard. 1969. Rabies epi- 
demiology in Ontario. Bulletin of the Wildlife Disease 
Association 5: 357-370. 

Kolenosky, G. B., D. R. Voigt, and R. O. Standfield. 
1978. Wolves and Coyotes in Ontario. Ontario Ministry 
of Natural Resources publication 18 pages. 

MacDonald, D. W., and D. R. Voigt. 1985. The biologi- 
cal basis of rabies models. Pages 71-108. in Population 
dynamics of rabies in wildlife. Edited by P. Bacon. 
Academic Press publishers. Toronto. 

Maclnnes, CD. 1987. Rabies. Pages 910-929 in Wild 
Furbearer Management and Conservation in North 
America. Edited by M. Novak, J. Baker, M. Obbard, and 
B. Malloch. Ontario Trappers Association Publishers, 
North Bay, Ontario. 

Rosatte, R. C, M. J. Power, and C. D. Maclnnes. 1991. 
Ecology of urban skunks, foxes and raccoons in 
Metropolitan Toronto. Pages 31-38 in Wildlife Conser- 
vation in Metropolitan Environments. Edited by L. W. 
Adams and D. L. Leedy. National Institute for Urban 
Wildlife Publishers, Columbia, Maryland. 

Rosatte, R. C, M. J. Power, C. D. Maclnnes, and J. B. 
Campbell. 1992. Trap-vaccinate-Release and oral vac- 
cination for rabies control in urban skunks, raccoons and 
foxes. Journal of Wildlife Diseases 28: 562-571. 

Rosatte, R. C, C. D. Maclnnes, M. J. Power, D. H. 
Johnston, P. Bachmann, C. P. Nunan, C. Wannop, 
M. Pedde, and L. Calder. 1993. Tactics for the control 
of wildlife rabies in Ontario Canada. Scientific and 
Technical Reviews of the Office of International Epi- 
zootics 12(1): 95-98. 

Rosatte, R. C, C. D. Maclnnes, R. Taylor Williams, and 
O. Williams. 1997. A proactive prevention strategy for 
raccoon rabies in Ontario, Canada. Wildlife Society 
Bulletin 25: 110-116. 

Tabel, H., H. Corner, W. Webster, and C. Casey. 1974. 
History and epizootiology of rabies in Canada. Canadian 
Veterinary Journal 15: 271-281. 

Voigt, D. R. 1987. Red Fox. Pages 379-392 in Wild 
Furbearer Management and Conservation in North 
America. Edited by M. Novak, J. Baker, M. Obbard, and 
B. Malloch. Ontario Trappers Association Publishers, 
North Bay, Ontario. 

Voigt, D. R., and D. W. MacDonald. 1984. Variation in 
the spatial and social behaviour of the red fox, Vulpes 
vulpes). Acta Zoologica Fennica 171: 261-265. 

Voigt, D. R., and W. E. Berg. 1987. Coyote. Pages 
345-357 in Wild Furbearer Management and Conser- 
vation in North America. Edited by M. Novak, J. Baker, 
M. Obbard, and B. Malloch. Ontario Trappers Associ- 
ation Publishers, North Bay, Ontaiio. 

Received 14 August 2000 
Accepted 20 March 2002 



132 



The Canadian Field-Naturalisx 



Vol. 116 



Predation by Wolves, Canis lupus, on Wolverines, Gulo gulo, and 
an American Marten, Martes americana, in Alaska. 



Kevin S. White'- 4, Howard N. Golden', Kris J. Hundertmark^, and Gerald R. Lee^ 

'Alaska Department of Fish and Game, Division of Wildlife Conservation, 333 Raspberry Road, Anchorage, Alaska 99518 

USA 
2 Alaska Department of Fish and Game, Division of Wildlife Conservation, 43961 Kalifomsky Beach Road, Suite B, 

Soldotna, Alaska 43961 USA 
3Basin Airmotive, P.O. Box 148, Glennallen, Alaska 99588 
^Present address: Alaska Department of Fish and Game, Division of Wildlife Conservation, PO Box 240020, Douglas, 

Alaska 99824, U.S.A. (e-mail:Kevin_white@fishgame. state. ak. us) 

White, Kevin S., Howard N. Golden, Kris J. Hundertmark, and Gerald R. Lee. 2002. Predation by Wolves, Canis lupus, on 
Wolverines, Gulo gulo, and an American Marten, Martes americana, in Alaska. Canadian Field-Naturalist 116(1): 
132-134. 

We report three instances of Wolf predation on mustelids in Alaska; two involved Wolverines and another involved an 
American Marten. Such observations are rare and in previous studies usually have been documented indirectly. This 
account provides insight into the potential role of Wolves in influencing mesocamivore conrmiunities in northern environ- 
ments. 

Key Words: Wolf, Canis lupus. Wolverine, Gulo gulo. Marten, Martes americana, predation, Alaska. 



Observations of Wolf (Canus lupus) -carnivore inter- 
actions are rare and have generally focused on those 
concerning ursids (Ursus arctos. Grizzly Bear; and 
U. americana. Black Bear; Murie 1944; Mech 1970, 
Rogers and Mech 1981; Ballard 1982) and canids 
{Canis latrans, Coyotes, and Vulpes vulpes. Red 
Foxes; Stenlund 1955; Mech 1970; Berg and 
Chesness 1978; Peterson 1996; Crabtree and 
Sheldon 1999). Nevertheless, direct and indirect evi- 
dence suggests that interactions involving actual and 
attempted Wolf predation on mustelids (Stenlund 
1955, Boles 1977, Route and Peterson 1991, Boyd et 
al. 1994, Paragi et al. 1996, Kohira and Rextad 1997; 
Mech et al. 1998) can occur at low frequency in 
some regions of North America and, further, may 
have a significant effect on mustelid populations 
(Palomares and Caro 1999). We report here on 
observations of Wolverine, Gulo gulo, and American 
Marten, Martes americana, predation by Wolves in 
Alaska. 

Account 1: On 20 March 1983 (1330 PST), during 
an aerial sex and age composition survey for Moose 
{Alces alces), we observed an adult Wolverine in the 
top of a large cottonwood {Populus trichocarpa) tree 
along the Chilkat River near Haines in southeastern 
Alaska (59° 37'N, 135° 55'W). Near the base of the 
tree was a pack of at least five Wolves in an area of 
blood-stained snow where the Wolves appeared to 
have been digging. One Wolf was observed with a 
juvenile Wolverine in its mouth. A few of the other 
Wolves were digging in the snow at what we sus- 
pected was a Wolverine den site, while another Wolf 
was lying down a short distance from the activity. 
Our impression was that the Wolves found the 
Wolverine den by chance and were in the process of 



digging it out to get to the kits. We were unable to 
determine if the Wolves actually consumed any of 
the kits, but judging from the extent and dispersion 
of blood on the snow, more than one kit had been 
killed. 

Account 2: We believe Wolves were responsible for 
the death of a yearling female Wolverine in an area 
of light spruce (Picea spp.) forest and tundra vegeta- 
tion in the Nelchina Basin in southcentral Alaska 
(62° 41'N, 147° 45'W). On 1 June 1997, while radio- 
tracking Wolverines as part of a population ecology 
study (Golden 1997), we observed a Wolf circling a 
yearling female Wolverine whose radio collar was 
on mortality mode and which showed no sign of 
movement (S. D. Bowen, Alaska Department of Fish 
and Game, personal communication). This was the 
first day we had detected her radio signal on mortali- 
ty mode since she was last seen alive on 15 May 
1997. We retrieved the carcass by helicopter on 2 
June 1997. While the carcass was being loaded onto 
the helicopter, two Wolves stood within 300 m bark- 
ing and howling (J. W. Testa, Alaska Department of 
Fish and Game, personal communication). The 
necropsy revealed five puncture holes in the skin, 
three in the chest and two in the groin, which may 
have been made by canine teeth of Wolves. Its chest 
was crushed laterally on the ventral side, resulting in 
several broken ribs. Although the carcass was in an 
advanced state of decomposition, it was intact and 
no part of it had been consumed. These observations 
plus the behavior of the Wolves and the timing of the 
death in late May suggest Wolves attacked and killed 
the Wolverine, possibly in defense of a wolf den site. 

Account 3: On 8 June 2000 (1222 ADT) during a 
telemetry re-location flight, we observed a radio- 



2002 



Notes 



133 



collared, solitary female Wolf in mixed- spruce {P. 
glauca and P. mariana) forest vigorously digging in 
moss-covered, hummocky soil, near Old Man Lake, 
Alaska (62° 31'N, 146° 81'W). As we circled the 
Wolf, we noticed one American Marten escaping 
through the forest -15 m away as the Wolf contin- 
ued digging in the original location. Subsequently, 
after ~2 minutes of digging, we observed the Wolf 
drag another Marten with its jaws from the under- 
ground cavity that it had been excavating. The Wolf 
then repeatedly bit, shook and dropped the Marten 
4-5 times until the Marten stopped moving, at which 
time it was presumed dead. The Wolf first stood 
guarding, and then rolled on the carcass until we ter- 
minated our observation at 1231 (ADT). 

Our observations indicated, to the extent possible, 
that Wolves did not feed on the carcasses of the 
Wolverines they had killed, a behavior noted by oth- 
ers (Burkholder 1962, Boles 1977). We can only 
speculate on the basis of that behavior, but ultimate 
explanations might include: elimination of competi- 
tors, defense of offspring, availability of prey, or dis- 
turbance by human observers. 

The role of Wolves in structuring mesocamivore 
communities is suspected to be significant though 
not fully understood. In areas where Wolves are re- 
colonizing historic ranges following prolonged 
absences, behavioral and ecological modifications of 
mesocarnivores can be dramatic (Crabtree and 
Sheldon 1999). Such changes underscore the impor- 
tance of Wolves in influencing ecosystem dynamics 
in such areas. While Wolf populations have fluctuat- 
ed historically in Alaska, their presence has 
remained constant and their role in influencing ungu- 
late populations has been studied extensively 
(Gasaway et al. 1983; Ballard et al. 1992; Dale et al. 
1994; Adams et al. 1995), though their interactions 
with mesocarnivores has received little attention. 
When considering species such as Wolverines that 
typically occur at low density, it is important to rec- 
ognize the role that even limited predation might 
exert on their population dynamics. Thus, in such 
cases, anecdotal accounts of predator- specific mor- 
tality provide valuable insights into the range of 
influence that Wolves might exert on mesocamivore 
communities in relatively undisturbed northern envi- 
ronments. 

Acknowledgments 

We thank S. D. Bowen, J. W. Testa, and D. 
Mortenson for their assistance with field observa- 
tions. We also thank S. Pyare and two anonymous 
reviewers for providing insightful comments on this 
manuscript. This work was funded by the Alaska 
Department of Fish and Game and Federal Aid to 
Wildlife Restoration Grant W-24-3, W-24-4, W-24- 
5,andW-27-l. 



Literature Cited 

Adams, L. G., B. W. Dale, and L. D. Mech. 1995. Wolf 
predation on caribou calves in Denali National Park, 
Alaska. Pages 245-260 in Ecology and conservation of 
wolves in a changing world. Edited by L. N. Carbyn, 
S. H. Fritts, and D. R. Seip. Canadian Circumpolar 
Institute, Occasional Publication Number 35. 

Ballard, W. B. 1982. Gray wolf-brown bear relationships 
in the Nelchina Basin of south-central Alaska. Pages 
71-80 in Wolves of the world: perspectives of behavior, 
ecology and conservation. Edited by F. H. Harrington, 
and P. C. Paquet. Noyes Publications, Park Ridge. New 
Jersey, USA. 

Ballard, W. B., J. S. Whitman, and D. J. Reed. 1991. 
Population dynamics of moose in south-central Alaska. 
Wildlife Monographs 114. 

Berg, W. E., and R. A. Chesness. 1978. Ecology of coy- 
otes in northern Minnesota. Pages 229-247 in Coyotes: 
biology, behavior, and management. Edited by M. 
Beckoff. Academic Press, New York, USA. 

Boles, B. K. 1977. Predation by wolves on wolverines. 
Canadian Field-NaturaUst 91: 68-69. 

Boyd, D. K., R. R. Ream, D. H. Fletcher, and M. W. 
Fairchild. 1994. Prey taken by colonizing wolves and 
hunters in the Glacier National Park area. Journal of 
Wildhfe Management 58: 289-295. 

Burkholder, B. L. 1962. Observations concerning wol- 
verine. Journal of Mammalogy 43: 263-264. 

Crabtree, R. L., and J. W. Sheldon. 1999. Coyotes and 
canid coexistence in Yellowstone. Pages 127-163 in 
Carnivores in ecosystems: the Yellowstone experience. 
Edited by T. W. Clark, A. P. Curlee, S. C. Minta. and 
P. M. Kareiva. Yale University Press, New Haven, 
Connecticut, USA. 

Dale, B. W., L. G. Adams, and R. T. Bowyer. 1994 
Functional response of wolves preying on barren-ground 
caribou in a multiple-prey ecosystem. Journal of Animal 
Ecology 63: 644-652. 

Gasaway, W. C, R. O. Stephenson, J. L. Davis, P. E. K. 
Shepherd, and O. E. Burris. 1983. Interrelationships 
of wolves, prey, and man in interior Alaska. Wildlife 
Monographs 84. 

Golden, H. N. 1997. Furbearer management techniques 
development: densides, trend, and harvest potential of 
wolverine populations. Alaska Department of Fish and 
Game, Federal Aid to Wildlife Restoration Research 
Grant W-24-5, Research Progress Report. 

Kohira M., and E. A. Rexstad. 1997. Diets of wolves, 
Canis lupus, in logged and unlogged forests of south- 
eastern Alaska. Canadian Field-Naturalist ill: 429-435. 

Mech, L. D. 1970. The wolf: ecology and behavior of an 
endangered species. Doubleday. New York. USA. 

Mech, L. D., L. G. Adams, T. J. Meier, J. W. Burch, and 
B. W. Dale. 1998. The wolves of Denali. University of 
Minnesota Press. Minneapolis, USA. 

Murie, A. 1944. The wolves of Mount McKinley. U.S. 
National Park Service. Fauna Series Number 5. 

Palomares, F., and T. M. Caro. 1999. Inlerspecitlc killing 
among mammalian carnivores. American Naturalist 153: 
492-508. 

Paragi, T. F., W. N. Johnson, D. D. Katnik, and A. J. 
Magoun. 1996. Marten selection of postfirc seres in 
the Alaskan taiga. Canadian Journal of Zoology 74: 
2226-2237. 



134 



The Canadian Field-Naturalist 



Vol. 116 



Peterson, R. O. 1996. Wolves as interspecific competi- 
tors in canid ecology. Pages 315-323 in Ecology and 
conservation of wolves in a changing world. Edited by 
L. N. Carbyn, S. H. Fritts, and D. R. Seip. Canadian 
Circumpolar Institute, Occasional Publication Number 
35. 

Rogers L. L., and L. D. Mech. 1981. Interactions of 
wolves and black bears in northeastern Minnesota. 
Journal of Mammalogy 62: 424-436. 

Route, W. T., and R. O. Peterson. 1991. An incident of 



wolf, Canis lupus, predation on a River Otter, Lutra 
canadensis, in Minnesota. Canadian Field-Naturalist 
105: 567-568. 
Stenlund, M. H. 1955. A field study of the timber wolf 
{Canis lupus) on the Superior National Forest, Min- 
nesota. Minnesota Department of Conservation, Tech- 
nical Bulletin Number. 4. 

Received 1 August 2000 
Accepted 25 March 2002 



Meek's Halfbeak, Hyporhamphus meeki, and Flying Gurnard, 
Dactylopterus volitans, captured in the Annapolis Basin, Nova Scotia 



A. Jamie F. Gibson^ and Ransom A. Myers^ 



'Acadia Centre for Estuarine Research, P.O. Box 115, Acadia University, Wolfville, Nova Scotia BOP 1X0 Canada; 

e-mail: jamie.gibson@acadiau.ca [Author to whom correspondence should be addressed] 
^Department of Biology, Dalhousie University, Halifax, Nova Scotia B3H 3J5 Canada; e-mail: myers@mscs.dal.ca 

Gibson, A. Jamie F., and Ransom A. Myers. 2002. Meek's Halfbeak, Hyporhamphus meeki, and Flying Gurnard 
Dactylopterus volitans, captured in the Annapolis Basin, Nova Scotia. Canadian Field-Naturalist 1 16(1): 134-135. 

We report the capture of several unusual fish at a hydroelectric generating station, Annapolis Royal, in the Annapolis 
Basin, Nova Scotia, during the fall of 1999. These include a Meek's Halfbeak, Hyporhamphus meeki, that is either the first 
or second Canadian record for this species and a Flying Gurnard, Dactylopterus volitans, that is a first record for the Bay 
of Fundy. Additionally, three Bluefish, Pomatomus saltatrix, and a Fourbeard Rockling, Enchelyopus cimbrius, were 
reported for the first time from the Annapolis Estuary during 1999. 

Key Words: Meek's Halfbeak, Hyporhamphus meeki. Flying Gurnard, Dactylopterus volitans, Bluefish, Pomatomus salta- 
trix, Fourbeard Rockling, Enchelyopus cimbrius, Annapolis River, Bay of Fundy, Nova Scotia, distribution. 



The Annapolis Basin in Nova Scotia is home to the 
western hemisphere's only tidal hydroelectric gener- 
ating station, located in Annapolis Royal (44° 45 'N, 
65° 3rW). As such, fish assemblages within the 
estuary are well studied, during both pre-operational 
stock assessments and surveys (e.g., Melvin et a. 
1985; Jessop 1976), and several assessments of fish 
passage facilities at the generating station (e.g., 
Gibson 1996). During 1999, several unusual fish 
specimens were captured while testing the effective- 
ness of an ultrasound fish diversion system at the sta- 
tion (Gibson and Myers 2000). A Meek's Halfbeak, 
Hyporhamphus meeki, captured on 23 September, 
1999, is either the first or second record of this 
species in Canada, and is the first record in 50 years. 
A Flying Gurnard, Dactylopterus volitans, captured 
28 September 1999, is a first record for the Bay of 
Fundy. Other unusual specimens captured at this 
location during September 1999 include three 
Bluefish, Pomatomus saltatrix and one Fourbeard 
rockling, Enchelyopus cimbrius. While these latter 
species occur regularly in the Bay of Fundy (Scott 
and Scott 1988), these species have not been reported 
previously in the Annapolis Estuary. 



Hyporhamphus spp. are planktivorous fish charac- 
terised by a very long lower jaw and short upper jaw. 
The taxonomic status of western Atlantic Hyporham- 
pus was clarified by the description of a new species, 
Hyporhamphus meeki, by Banford and Collette 
(1993). This species ranges north from the Gulf of 
Mexico, and is a rare stray into the Gulf of Maine. It 
usually can be distinguished from its southern rela- 
tive, Hyporhamphus unifasciatus, by having a greater 
number of total gill rakers on the first (3 1 to 40) and 
second arches (20 to 30), and having a ratio of preor- 
bital length to orbit diameter greater than 0.70 
(Banford and Collette 1993). Contreras-Balderas et al. 
(1997) suggest the species can be distinguished using 
the shape of the lateral band. The specimen reported 
herein is unusual in having 29 gill rakers on right first 
arch and 23 gill rakers on the right second arch, both 
of which are low for this species. The identifying 
characteristic for this specimen is the ratio of preor- 
bital length to orbit diameter, which equals to 0.74 
(Bruce Collette, personal communication). 

One previous record of Hyporhamphus sp. exists 
for eastern Canada. This specimen, reported as 
Hyporhamphus unifasciatus, was captured in a 



II 




2002 



Notes 



135 



herring weir at Chamcook, New Brunswick, on 30 
September, 1949 (Leim and Day 1959). Its location 
suggests that this specimen may actually have been a 
H. meeki, sl species that was not described at the time 
that this specimen was captured. 

Dactylopterus volitans is a benthic species of fish 
that superficially resembles a searobin, and is typically 
found in tropical or warm temperature water. It ranges 
south to Argentina, and commonly is found as far 
north as North Carolina. The species occasionally 
strays north to Nova Scotia. Scott and Scott (1988) 
report four specimens from the Atlantic coast of Nova 
Scotia during the 1970s. The Nova Scotia Museum of 
Natural History has two records of D. volitans in Nova 
Scotia, both from the Atlantic Coast in 1990 (John 
Gilhen, personal communication). D. volitans have 
also been reported from the Grand Banks and Scotian 
Shelf (Lou VanGuelpen, personal communication). 

Bluefish are regular summer visitors to the Bay of 
Fundy during warmer summers. The Fourbeard 
Rockling is a resident of the Bay of Fundy, and some- 
times moves into embayments during summer and fall 
(Scott and Scott 1988). Water temperature was warmer 
and salinity was higher during early September in 
1999 than during six previous years of samphng since 
1985, perhaps leading to favourable conditions for 
these fish to move farther into the estuary. 

The Meek's Halfbeak (reference number: ARC 
0016423) and Flying Gurnard (reference number: 
ARC 0016422) are deposited at the Atlantic 
Reference Centre located at the Huntsman Marine 
Science Centre, St. Andrews, New Brunswick. 

Acknowledgments 

The authors express their thanks to John Gilhen 
(Nova Scotia Museum of Natural History, 1747 
Summer Street, Halifax, Nova Scotia, B3H 3A6), 
Lou VanGuelpen (Atlantic Reference Centre, 
Huntsman Marine Science Centre, 1 Lower Campus 
Road, St. Andrews, New Brunswick, E5B 2L7), 
Donald F. McAlpine (New Brunswick Museum, 277 
Douglas Avenue, Saint John, New Brunswick, E2K 
1E5) for searching their collections for records of 
these species. Thanks are also extended to Bruce 
Collette (Systematics Laboratory, National Marine 
Fisheries Services, Smithonian Institution, 



Washington, DC, 20560) for assistance with the 
identification of the H. meeki. 

Literature Cited 

Banford, H. M., and B. B. CoUete. 1993. Hyporhamphus 
meeki, A new species of halfbeak (Teleostei: 
Hemiramphidae) from the Adantic and Gulf coasts of 
the United States. Proceedings of the Biological Society 
of Washington 106: 369-384. 

Contreras-Balderas, S., M. L. Lozano-Vilano, and M. E. 
Garcia Ramirez. 1997. Distributional and ecological 
notes on the halfbeaks of eastern Gulf of Mexico, with a 
provisional key for their identification. Gulf Research 
Reports 9: 327-331. 

Gibson, A. J. F. 1996. Distribution and seaward migra- 
tion of young-of-the-year American Shad {Alosa 
sapidissima), Blueback Herring (A. aestivalis) and 
Alewives (A. psuedoharengus) in the Annapolis River 
Estuary. M. Sc. thesis. Acadia University, Wolfville, 
Nova Scotia. 96 pages. 

Gibson, A. J. F., and R. A. Myers. 2000. Assessment of a 
high-frequency sound fish diversion system at the 
Annapolis Tidal Generating Station with notes about the 
survival of fish moving seaward at the Annapolis cause- 
way. Acadia Centre for Estuarine Research Publication 
Number 55. Acadia University, Wolfville, Nova Scotia. 
98 pages. 

Jessop, B. M. 1976. Physical and Biological Survey of 
the Annapolis River, 1975. Data Record Series Number 
MAR/D-76-8. Fisheries and Marine Service. Department 
of the Environment, Halifax. 29 pages. 

Leim, A. H., and L. R. Day. 1959. Records of uncommon 
and unusual fishes from eastern Canadian waters. 1950- 
1958. Journal of the Fisheries Research Board of Canada 
16:503-514. 

Melvin, G. D., M. J. Dadswell, and J. D. Martin. 1985. 
Impact of Lowhead Hydroelectric Tidal Power 
Development on Fisheries I. A Pre-operation Study of 
the Spawning Population of American Shad Alosa 
sapidissima (Pisces:Clupeidae). in the Annapolis River, 
Nova Scotia, Canada. Canadian Technical Report of 
Fisheries and Aquatic Sciences Number 1340. St. 
Andrews, New Brunswick. 33 pages. 

Scott, W. B., and M. G. Scott. 1988. Adantic Fishes of 
Canada. Canadian Bulletin of Fisheries and Aquatic 
Sciences 219. 731 pages. 

Received 5 June 2000 
Accepted 1 9 February 2002 



136 



The Canadian Field-Naturalist 



Vol. 116 



Apparent Capture Myopathy in Hoary Bats, Lasiurus cinereus: 
A Cautionary Note 



Thomas S. Jung'-2, Ian D. Thompson^, M. Brian C. Hickey^-^ and Rodger D. Titmani 

'McGill University, Department of Natural Resource Sciences, 21111 Lakeshore Blvd., Montreal, Quebec, H9X 3V9 

Canada 
2Present address: Yukon Department of Renewable Resources, Fish and Wildlife Branch, P.O. Box 2703, Whitehorse, 

Yukon, YIA 2C6 Canada; thomas.jung@gov.yk.ca 
3Canadian Forest Service, Great Lakes Forestry Centre, 1219 Queen Street East, Sault Ste. Marie, Ontario, P6A 5M7 

Canada 
^York University, Department of Biology, 4700 Keele St., North York, Ontario, M3J 1P3 Canada 
sPresent address: St. Lawrence River Institute of Environmental Sciences, Cornwall, Ontario, K6H lEl Canada 

Jung, Thomas S., Ian D. Thompson, M. Brian C. Hickey, and Roger D. Titman. 2002. Apparent capture myopathy in 
Hoary Bats, Lasiurus cinereus: a cautionary note. Canadian Field-NaturaUst 116(1): 136-137. 

Capture myopathy, a stress-induced disease resulting in metabolic acidosis, has not been recorded in bats. We observed 
two cases of apparent capture myopathy in Hoary Bats {Lasiurus cinereus) captured in mist-nets. We caution those intend- 
ing to work with this species of a potential occurrence, and discuss capture protocols that may reduce the risk of capture 
myopathy. 

Key Words: Hoary Bats, Lasiurus cinereus, capture myopathy, live-capture. 



Biologists often rely on data obtained through live- 
capture. Live-capturing animals, hov^ever, may result 
in capture myopathy: a stress-induced disease, where- 
by the liberation of large amounts of lactic acid 
results in metabolic acidosis, which may lead to mus- 
cle necrosis and death (Chalmers and Barrett 1982; 
Pond and O'Gara 1994). Clinical signs of this condi- 
tion may include muscle stiffness, loss of coordina- 
tion, listlessness, and increased respiration and body 
temperatures, but affected animals can die without 
exhibiting outward symptoms (Chalmers and Barrett 
1982). Capture myopathy can occur immediately 
upon capture or up to one month afterwards, and has 
been reported from a variety of species. To the best 
of our knowledge, there are no published reports of 
bats succumbing to capture myopathy. We encourage 
others to collate and disseminate their observations of 
capture myopathy in bats. 

As part of a study on the habitat ecology of forest- 
dwelling bats in central Ontario (46.8° N; 81.9° W; 
Jung et al. 1999), bats were live-captured in mist- 
nets set across rivers and logging roads. On 4 August 
1995 at 22:40 (EDT) an adult male Hoary Bat was 
captured. The bat was extracted from the net in <5 
min, and was active and vocal when captured. Once 
extracted from the net, the bat was placed in a home- 
made steel mesh holding container (Kunz and Kurta 
1988), for about 25 min. After which the bat was dis- 
covered languid and did not fly. Therefore, it was 
placed on the limb of a small Trembling Aspen 
(Populus tremuloides), where it remained, suspended 
by its hind feet. The next morning the bat was dis- 
covered dead. Post-mortem examination indicated 
that the bat was in good condition, as the pelage was 
shiny, there were few noticeable ectoparasites, and 
its mass appeared normal (29.8 g.). Due to the diffi- 



culty in keeping the specimen frozen at a remote 
field camp, it was not sent for necropsy. We do not 
believe that the Hoary Bat was excessively handled, 
relative to > 100 Myotis spp. that we had captured. It 
is our belief that the Hoary Bat died as a result of the 
stress of being captured. 

Similarly, as part of a study of the thermal ecology 
and foraging behavior of female Hoary Bats in 
southwestern Ontario (43.3° N; 81.8° W; Hickey and 
Fenton 1996), Hoary Bats were live-captured near 
street lights by swinging mist-nets at diving bats. On 
8 June 1989, an adult female Hoary Bat was cap- 
tured and banded. After about 10 min the bat was 
hand-released, but was listless and did not fly away. 
The bat was placed on the limb of a nearby tree and 
it remained there. The bat was checked daily and had 
not moved. On 12 June 1989, the bat was discovered 
dead, having apparently not moved since being cap- 
tured. The bat did not appear injured, and seemed to 
have perished as a result of capture myopathy. In a 
related manner, our experience (MBCH) with Hoary 
Bats in captivity suggests that these species have a 
propensity to die within 1 month of capture, which 
may also be stress-related. 

This is the first known report of apparent stress- 
induced capture myopathy in bats. We do not know 
why Hoary Bats may be more susceptible to capture 
myopathy than other bats such as Myotis spp. 
Although the prognosis is poor for individuals exhibit- 
ing signs of capture myopathy. Pond and O'Gara 
(1994) state that keeping the animal well-oxygenated 
and providing intravenous or subcutaneous adminis- 
tration of sodium bicarbonate in a saline solution is a 
recommended therapy. If stress-induced capture 
myopathy does occur somewhat regularly in Hoary 
Bats, then the use of a chemical immobilization agent 



2002 



Notes 



137 



may reduce the likelihood of capture myopathy, 
through a reduction in the onset of extreme stress. 
Chemical tranquilizers have been used for small 
mammals (e.g., Wright 1983), but are not commonly 
used on bats. We recommend that Hoary Bats be pro- 
cessed as quickly as possible after live-capture, as 
they may not be as resilient to capture as other species 
of bats. We also suggest that steel mesh containers 
may provide substantially more oxygen than cloth 
bags, which are sometimes used to hold bats, thereby 
reducing the likehhood of capture myopathy. Bats that 
do perish after capture should be sent for necropsy to 
verify the cause of death. 

Acknowledgments 

R. M. Brigham provided thoughtful comments on 
an earlier draft of this note. We thank the field assis- 
tants that participated in mist-netting. Live-capture 
of bats was conducted under the approval of animal 
care committees at McGill and York universities. 

Literature Cited 

Chalmers, G. A., and M. W. Barrett. 1982. Capture 
myopathy. Pages 84-94. In Noninfectious diseases of 



wildlife. Edited by G. L. Hoff and J. W. Davis. Iowa State 
University Press. Ames, Iowa. 174 pages. 

Hickey, M. B. C, and M. B. Fenton. 1996. Behavioural 
and thermoregulatory responses of female Hoary Bats, 
Lasiurus cinereus, to variations in prey availability. 
Ecoscience 3: 412-422. 

Jung, T. S., I. D. Thompson, R. D. Titman, and A. P. 
Applejohn. 1999. Habitat selection by forest-dwelling 
bats in relation to stand type and structure in central 
Ontario. Journal of Wildlife Management 63: 1306-1319. 

Kunz, T. H., and A. Kurta. 1988. Capture methods and 
holding devices. Pages 1-30 In Ecological and behav- 
ioral methods for the study of bats. Edited by T. H. 
Kunz. Smithsonian Institution Press, Washington, D.C. 

Pond, D. B., and B. W. O'Gara. 1994. Chemical immobi- 
lization of large mammals. Pages 125-139 In Research 
and management techniques for wildlife and habitats. 
Fifth ed. Edited by T. A. Bookhout. The Wildlife Society, 
Bethesda, Maryland. 740 pages. 

Wright, J. M. 1983. Ketamine hydrochloride as a chemi- 
cal restraint for selected small mammals. Wildlife 
Society Bulletin 11:76-79. 

Recieved 9 June 2000 
Accepted 22 March 2002 



Mobbing Black-billed Magpie, Pica hudsonia. Killed by 
Cooper's Hawk, Accipiter cooperii 

Geoffrey L. Holroyd 

Canadian Wildlife Service, Room 200, 4999-98 Avenue, Edmonton, Alberta T6B 2X3 Canada 

Holroyd, Geoffrey L. 2002. Mobbing Black-billed Magpie, Pica pica, killed by Cooper's Hawk, Accipiter cooperii. 
Canadian Field-Naturahst 116(1): 137-138. 

A Cooper's Hawk killed a Black-billed Magpie that was part of a group mobbing it. This occurrence happened before the 
breeding season. Any benefit of mobbing did not enhance the survival of this magpie. 

Key Words: Black-billed Magpie, Pica hudsonia. Cooper's Hawk, Accipiter cooperii, mobbing, mortality. 



Mobbing is the massing together of birds, often of 
different species, for the purpose of aggression and 
attack against a common predator (Terres 1982). 
Mobbing is most prevalent in the nesting season 
(Shedd 1 982) but does occur year-round, particularly 
by species that remain on their nesting territory 
(Shedd 1983). This note documents an observation of 
a Cooper's Hawk (Accipiter cooperii) killing and eat- 
ing a Black-billed Magpie (Pica hudsonia) that was 
one of a group that was mobbing it. 

On 15 April 2000, just after 17:00 h, Phil Trefry 
drew our attention to a Cooper's Hawk on a White 
Birch (Betula papyrifera) in a low lying birch-willow 
(Salix spp.) bluff in a depression in front of his home 
about 50 km east of Edmonton (54° 24'N 1 12° 
50'W). The farm is in rolling aspen parkland with 
open fields and Trembling Aspen (Popuhts tremu- 



loides) and Balsam Poplar (P. balsamifera) dominat- 
ed woods. The hawk was being mobbed by four mag- 
pies and two Blue Jays (Cyanocitta cristata). The 
mobbing was mostly "vocal approach" (sensu Shedd 
1982) and flying at a distance. The mobbers did not 
fly at or strike the hawk. A Red-tailed Hawk (Biiteo 
jamaicensis) was perched 50 m west of the Cooper's 
Hawk, higher in a birch tree, but was ignored by the 
corvids. One agitated magpie perched 1-2 m above 
the Cooper's Hawk in the same tree, calling and hop- 
ping ("mobbing" sensu Shedd 1982). The hawk glid- 
ed to the ground under overhanging willow branches, 
10 m to the south of its original perch. The magpie 
hopped to the ground about 3 m from the hawk where 
they were hidden from the observers by tall dead 
herbaceous vegetation. The hawk flew to the magpie, 
killed it and carried it aloim the edue o{ the bluff. 



138 



The Canadian Field-Naturalist 



Vol. 116 



50 m north, then into the bluff behind some birch trees 
and out of sight. Over 10 magpies and two Blue Jays 
arrived from different directions and they and the 
other corvids called, hopped agitatedly and flew over- 
head. The Red-tailed Hawk and another that had been 
perched to the east, flew closer and perched nearby. 
After about 10 minutes the corvids and red-tails left. 
Three hours later we visited the two sites. At the kill 
site we could see the wing-tip prints of the Cooper's 
Hawk in the snow and two marks between the prints, 
presumably where the magpie had landed and died. At 
the northerly destination was a collection of magpie 
body feathers over an area of 3 sq. m, where the hawk 
had plucked part of the magpie. There were no flight 
feathers nor any of the magpie's carcass. 

The purpose of mobbing is to drive a predator from 
a nearby nest or territory (Shedd 1982). Mobbing 
birds have been observed driving aerial predators to 
the ground or into the water (Terres 1982: 63, 627). 
However, mobbing does not always drive a predator 
from a nest (Taylor 1972). The present observation is 
unusual because the magpie was killed while mob- 
bing and magpies are rarely preyed upon by raptors. 

In April, Black-billed Magpies can be expected on 
territory. Although no magpies nested in the bluff 
where this observation took place, they are common 
breeding birds in the area (personal observation). 
Mobbing is more common in the breeding season in 
migratory species such as the American Robin 
(Shedd 1982), but occurs year-round with no appar- 
ent seasonality in non-migratory species such as 
Black-capped Chickadee {Poecile atricapillus) 
(Shedd 1983). The presumed advantage of mobbing 
to a bird without a nest is to drive the potential 
predator from the permanent territory of the bird. In 
our observation, the magpie did not have an active 
nest in the immediate vicinity to protect. 

Sordahl (1990) reviewed 30 instances of predators 
killing mobbing birds. At least one of these instances 
involved a Cooper's Hawk (Wilson 1986). In 1962 
in Virginia, R. D. Denson (1979) observed a Great 
Horned Owl {Bubo virginianus) kill one of a flock of 
American Crows {Corvus brachyrhychos) that were 
mobbing it. The owl grabbed the crow out of the air 
from its perch. As in our observation, the crow went 
too close to the owl. Although mobbing behaviour 
may drive away a predator, the close proximity of 
predator and potential prey can be expected to have 
some risk for the prey. In this case the magpie 
appeared to have made a mistake when it landed on 
the ground too near the hawk. The magpie was killed 
with little effort by the Cooper's Hawk as evidence 
by the lack of sign of a struggle in the snow. 

Passerines mobbed Northern Harriers {Circus cya- 
neus) only when the harriers are in flight and active- 
ly hunting (Bildstein 1982). When the harriers are 
perched, the passerines ignore the hawks. In both our 
observation and that of Denson (1979) the Great 
Horned Owl and the Cooper's Hawk, species that 
will hunt from a perch, were mobbed while perched. 



The nearby perched red-tails, which are a known 
nest predator (Trost 1999), were not mobbed. 

Magpies are rarely taken by raptors (Trost 1999). 
Three magpies were recorded in 550 prey remains 
collected at Peregrine Falcon {Falco peregrinus) 
nests (unpublished data) but none in 203 prey at 
Prairie Falcon {F. mexicanus) nests in southern 
Alberta (Hunt 1983). One Cooper's Hawk nest 
observed at nearby Beaverhill Lake did not include 
any magpie remains (unpublished data). Magpies 
were not reported in the diet of Cooper's Hawks in 
Washington and Oregon, which are within the mag- 
pie's range (Kennedy and Wilson 1986; Reynolds 
and Meslow 1984). Although this observation is only 
of a single event, the killing of a mobbing magpie by 
a Cooper's Hawk has interesting implications. 

Acknowledgments 

I thank Elisabeth Beaubien, Martin Raillard, Tony 
Erskine and an anonymous reviewer for comments 
on earlier draft of this manuscript. Elisabeth, Helen 
Trefry and Phil Trefry also observed this event. 

Literature Cited 

Bildstein, K. L. 1982. Responses of Northern Harriers 
to mobbing passerines. Journal of Field Ornithology 
53: 7-14. 

Denson, R. D. 1979. Owl predation on a mobbing crow. 
Wilson Bulletin 91: 133. 

Hunt, L. E. 1983. Diet and habitat use of nesting Prairie 
Falcons {Falco mexicanus) in an agricultural landscape 
in southern Alberta. MSc. Thesis, University of Alberta. 
60 pages. 

Kennedy, P. L., and D. R. Wilson. 1986. Prey size selec- 
tion in nesting male and female Cooper's Hawks. Wilson 
Bulletin 98: 110-115. 

Reynolds, R. T., and E. C. Meslow. 1984. Partitioning of 
food and niche characteristics of coexisting Accipiter 
during breeding. Auk 101: 761-779. 

Shedd, D. H. 1982. Seasonal variation and function of 
mobbing and related behaviors of the American Robin 
{Turdus migratorius). Auk 99: 342-346. 

Shedd, D. H. 1983. Seasonal variation in mobbing inten- 
sity in the Black-capped Chickadee. Wilson Bulletin 95: 
343-348. 

Sordahl, T. A. 1990. The risks of avian mobbing and dis- 
traction behaviour: an anecdotal review. Wilson Bulletin 
102: 349-352. 

Taylor, W. K. 1972. Mobbing of a Fish Crow by passe- 
rines. Wilson Bulletin 84: 98. 

Terres, J. K. 1982. The Audubon Society Encyclopedia of 
North American Birds. Knopf, New York. 1 109 pages. 

Trost, C. H. 1999. Black-billed Magpie {Pica pica). In 
The Birds of North America, Number 389. Edited by A. 
Poole and F. Gill. The Birds of North America, Inc., 
Philadelphia, Pennsylvania. 28 pages. 

Wilson, R. E. 1986. An aerial counter attack by a 
Cooper's Hawk. Bulletin of the Texas Ornithological 
Society 19:32-33. 

Received 21 September 2000 
Accepted 18 March 2002 



2002 



Notes 



139 



Breeding Season of Wolves, Canis lupus, in Relation to Latitude 

L. David Mech 

Biological Resources Division, U.S. Geological Survey, Northern Prairie Wildlife Research Center, 8711-37th Street, SB, 

Jamestown, North Dakota 58401-7317 
•Mailing address: The Raptor Centre, University of Minnesota, 1920 Fitch Avenue, St. Paul, Minnesota 55108 USA 

Mech, L. David. 2002. Breeding season of Wolves, Canis lupus, in relation to latitude. Canadian Field Naturalist 116(1): 
139-140. 

A significant relationship was found between Wolf {Canis lupus) breeding dates and latitudes between 12° and 80° N, with 
Wolves breeding earlier at lower latitudes, probably because of differences in seasonality. 

Key Words: Wolf, Canis lupus, reproduction, latitude, breeding, mating. 



A general relationship between breeding dates in 
Wolves {Canis lupus) and latitudes from 41°-71°N 
was noticed by Mech (1970: 117) when he summa- 
rized published data from several locales. However, 
he conducted no statistical test of this hypothesis. 
Herein, I add data from other areas of latitude from 
12° to 80° N (Table 1) and statistically test the effect 
of latitude. 

I used latitude as the independent variable in a sim- 
ple linear regression and the reported breeding date as 
the dependent variable. For breeding date, I used the 
mid date for the reported breeding season and con- 
verted all dates to sequential numbers starting with 
15 October to facilitate comparing breeding dates 
before and after the start of the calendar year. 

The relationship between breeding date and lati- 
tude was highly significant (r2=0.74; P< 0.0001; 
y = 16.19 + 2.23x), supporting Mech's (1970) 
hypothesis. On average, breeding season shifts 22 



days later with each 10° latitude increase. It seems 
reasonable to suggest that the shift is related to differ- 
ences in general seasonality and thus in associated 
ecological conditions. 

Acknowledgments 

This study was supported by the Biological 
Resources Division of U.S. Geological Survey and 
the U.S. Department of Agriculture, North Central 
Research Station. 

Literature Cited 

Bailey, V. 1926. A biological survey of North Dakota. U.S. 
Department of Agriculture, Biological Survey, North 
American Fauna 49. 

Cowan, I. M. 1947. The timber wolf in the Rocky 
Mountain national parks of Canada. Canadian Journal 
Research 25: 139-174. 

Fritts, S. H., and L. D. Mech. 1981. Dynamics, move- 
ments, and feeding ecology of a newly protected wolf 



Table 1 . Mating seasons of wolves at various latitudes. 



Location 


N Latitude 


Season 


Authority 


Southern India 


12° 


October 


Kumar and Rahmani 2001 


Arizona^ 


34° 


February, March 


W. Brown, personal communication 


Illinois^ 


42° 


February 


Rabb 1968 


Yellowstone National Park'' 


45° 


February 


Smith etal. 1998 


Ontario 


47° 


Early March 


Joslin 1966 


North Dakota 


46°^9° 


January 


Bailey 1926 


Isle Royale (Michigan) 


47° 


Late February 


Mech 1966 


Minnesota 


48° 


February 


Mech and Knick 1978; Fritts and Mech 
1981; Fuller 1989 


British Columbia 


5r-53° 


March 


Cowan 1947 


Germany^ 


52° 


Mid March 


Schonberner 1965 


Alberta 


60° 


February. March 


Soper 1942; Fuller and Novakowski 1955 


Northwest Territories 


60°-65° 


Late March 


Kelsall 1960 


Alaska 


60°-7r 


March 


Murie 1944; Kelly 1954; Rausch 1967;, 
Lentfer and Sanders 1973; 
Mech etal. 1998 


Finland 


60°-70 


March 


Pulliainen 1965 


Russia 


71° 


Late March-early April 


Makridin 1962 


Ellesmere Island 


80° 


Late March-early April 


Mech 1993 



''Captive wolves 
Wolves translocated from 53-56° N. 



140 



The Canadian Field-Naturalist 



Vol. 116 



population in northwestern Minnesota. Wildlife Mono- 
graph 80. 

Fuller, T. K. 1989. Population dynamics of wolves in 
north-central Minnesota. Wildlife Monograph 105. 

Fuller, W. A., and N. S. Novakowski. 1955. Wolf control 
operations, Wood Buffalo National Park, 1951-1952. 
Canadian Wildlife Service, Wildlife Management Bul- 
letin Series 1, Number 1 1. 

Joslin, P. W. B. 1966. Summer activities of two timber 
wolf {Canis lupus) packs in Algonquin Park. M.S. the- 
sis. University of Toronto, 99 pages. 

Kelly, M. W. 1954. Observations afield on Alaskan 
wolves. Proceedings of Alaska Science Conference 5: 35. 

Kelsall, J. P. 1960. Co-operative studies on barren ground 
caribou 1957-58. Canadian Wildlife Service, Wildlife 
Management Bulletin Series 1, Number 12. 

Kumar, S., and A. R. Rahmani. 2001. Livestock depre- 
dation by wolves in the great Indian Bustard Sanctuary, 
Nannaj (Maharashtra), India. Journal, Bombay Natural 
History Society 97: 340-348. 

Lentfer, J. W., and D. K. Sanders. 1973. Notes on the 
captive wolf {Canis lupus) colony, Barrow, Alaska. 
Canadian Journal of Zoology 5 1 : 623-627. 

Makridin, V. P. 1962. The wolf in the Yamal north. 
Zoological Zhumal 41: I4I3-I4I7. 

Mech, L. D. 1966. The wolves of Isle Royale. U. S. 
National Park Service Fauna Series Number 7. 

Mech, L. D. 1970. The wolf: ecology and behavior of an 
endangered species. Natural History Press, Doubleday 
Publishing Co., New York. 



Mech, L. D. 1993. Resistance of young wolf pups to 

inclement weather. Journal of Mammalogy 74: 485-^86. 
Mech, L. D., and S. T. Knick. 1978. Sleeping distances in 

wolf pairs in relation to breeding season. Behavioral 

Biology 23: 521-525. 
Mech, L. D., L. G. Adams, T. J. Meier, J. W. Burch, and 

B. W. Dale. 1998. The wolves of Denali. University of 

Minnesota Press, Minneapolis. 
Murie, A. 1944. The wolves of Mount McKinley. U.S. 

National Park Service Fauna Series Number 5. 
Pulliainen, E. 1965. Studies of the wolf {Canis lupus L.) 

in Finland. Annales Zoologici Fennici 2: 215-259. 
Rabb, G. B., J. H. Woolpy, and B. E. Ginsburg. 1967. 

Social relationships in a group of captive wolves. Ameri- 
can Zoologist 7: 305-312. 
Rausch, R. A. 1967. Some aspects of the population ecol- 
ogy of wolves, Alaska. American Zoologist 7: 253-265. 
Schonberner, D. 1965. Observations on the reproductive 

biology of the wolf. Zeitschrift fur Sauzetierkunde 30: 

171-178. 
Smith, D. W., K. M. Murphy, and D. S. Guernsey. 1998. 

Yellowstone Wolf Project. Annual Report YCR-NR-99-1. 

Yellowstone Center for Resources, Yellowstone National 

Park, Wyoming. 
Soper, J. D. 1942. Mammals of Wood Buffalo Park, 

northern Alberta and District of Mackenzie. Journal of 

Mammalogy 23: 119-145. 

Received 2 October 2000 
Accepted 5 February 2002 



Canada and the "Buffalo", Bison bison: A Tale of Two Herds 

W. A. Fuller 

Professor Emeritus, University of Alberta; Adjunct Professor Athabasca University, Box 672, Athabasca, Alberta T9S 2A6 

Fuller, W. A. 2002. Canada and the "buffalo". Bison bison: A tale of two herds. Canadian Field-Naturalist 116(1): 
141-159. 

From 1907 to 1912 the Canadian government purchased and imported more than 700 plains Bison, Bison bison, from 
Michel Pablo in Montana. A new national park, with an area of 159 square miles was established near Wainwright, 
Alberta, to accommodate them. It has generally been acknowledged that Buffalo National Park played an important role in 
saving the Plains Bison from extinction. This paper makes use of a packet of government files that were saved from 
destruction during the early 1940s. The files deal mainly with events from 1912 to 1925, including the first appearance of 
bovine tuberculosis, and later the prevalence of tuberculosis in the herd. They also contain notes from the meetings of 
senior civil servants that led to the decision to transfer diseased plains bison to Wood Buffalo National Park, as well as 
summaries of submissions of those opposed to the transfer. One option, to slaughter the entire herd and start over with dis- 
ease-free stock, was rejected by well-meaning members of the public. When the Buffalo National Park was turned over to 
the military in 1940, 17 000 bison had been slaughtered as a result of annual culling. Ironically, had a total slaughter been 
carried out in 1923, fewer than 7 000 would have been killed. In addition, it is probable that we would have pure Wood 
Bison and no tuberculosis in Wood Buffalo National Park. In 1963, 18 disease-free Bison derived from a group of animals 
that showed some of the characteristics of Wood Bison, were released in the Mackenzie Bison Sanctuary. That herd now 
numbers about 2 600 individuals. As in 1923 we again have two herds, one with a high prevalence of tuberculosis and a 
second that is disease-free. In 1990 an Environment Panel (1990) recommended total depopulation of Wood Buffalo 
National Park and restocking with disease-free animals. As in 1923 the recommendation to slaughter and restock met oppo- 
sition on several fronts and so far no action has been taken. Must we repeat the serious error made in 1923? 

Key Words: Wainwright, Bison, tuberculosis, brucellosis, Pablo, slaughters, cattalo, transfer. 



More than thirty years ago I received from Dr. 
C. H. Douglas Clarke (see tribute and bibliography 
by Lumsden 1984a, 1984b) a box of old federal gov- 
ernment files that were related to the origin and sub- 
sequent operation of the Buffalo National Park near 
Wainwright, Alberta, and the transfer of Bison from 
there to Wood Buffalo National Park. According to a 
letter that accompanied the files, they were "...sal- 
vaged from the wartime file disposal in Ottawa." 
Clarke gave as his reason for saving the files from 
destruction that "I realized that it was a personal 
record, kept by the late Maxwell Graham, to show 
that for his part he did his best to prevent the transfer 
of buffalo from Wainwright to the Wood Buffalo 
Park, and to promote herd reduction by slaughter." 

Maxwell Graham was Director of Animals in the 
Dominion Parks Branch of the Department of the 
Interior from January 1912 until his transfer to a dif- 
ferent Branch on 1 January 1922. The files shed light 
on his role as guardian of the Bison in the Buffalo 
National Park near Wainwright, Alberta. At one 
time, those buffalo were much in the news, and some 
of the material in the files would have been "hot 
stuff politically. Clarke wrote: "Poor old Graham 
was in an uncomfortable spot, and I have a feeling of 
personal obligation towards him." By accepting the 
files, I assumed that obligation, which I am only now 
attempting to fulfill. 

This paper will reveal information that has not 
been made public until now about the management 



of Buffalo National Park and about the transfer of 
buffaloes to the newly created Wood Buffalo 
National Park. My belief is that lessons learned so 
long ago are pertinent to the current situation in 
Wood Buffalo National Park. My hope is that those 
lessons will be taken to heart by all who have an 
interest in the long-term welfare of the buffalo.The 
files are the source of all direct citations set in small 
type and inset paragraphs. 

Plains Buffalo 

1. Disappearance from the wild 

When Europeans first reached the great central 
plain of North America they were greeted by 
immense herds of "buffalo", whose Latin name is 
Bison bison. Bison is now the approved common 
name for the beast, and I will use it from this point 
on. As time wore on and the human population 
increased, the Bison population decreased. The story 
of their decline has been told by several observers, 
such as Allen (1875), Hornaday (1889) and Isenberg 
(2000) for the United States; Hewitt (1921). Roe 
(1951) and Foster (1992) for Canada. Around the 
middle of the 19th century so many Bison had been 
killed that those remaining were essentially confined 
to two herds that were referred to as "southern" and 
"northern." The southern herd was wiped out during 
the 1870s, and the northern herd followed in the 
188()s. Fortunately, a number of ranchers, in both the 
United States and Canada, had captured and raised a 



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few Bison. One such herd is referred to either as the 
Allard, or the Pablo, herd. That herd was in 
Montana, conveniently close to Canada. 

2. Recovery in Buffalo National Park 

After a considerable amount of negotiation, the 
Canadian government reached an agreement in 1907 
with the owners of the Pablo herd to purchase a few 
hundred bison at $200 per head. They were to be 
delivered to a new park close to Wainwright, 
Alberta. However, the first shipment arrived before 
the fencing and buildings were finished in the new 
park, and the animals had to be released in Elk Island 
National Park, which had been set aside for the 
preservation of "elk" (American Elk, Cervus 
elaphus) in 1904. 

When Pablo began rounding up his animals, he 
discovered that he had more than the few hundred 
bison that Canada had originally contracted for. On 
11 June 1907, 199 head arrived at Lamont, Alberta, 
the rail station nearest to Elk Island National Park. 
On 11 October, another shipment of 211 head 
arrived at Lamont. No shipments were made during 
1908 owing to illness on the part of Pablo, and the 
escape of 120 animals ready for transfer, but, during 
that year 73 miles (1 16 km) of fencing was put in 
place, and construction of other facilities was com- 
pleted at the new Buffalo National Park. In May of 
1909, 325 of the Bison originally sent to Elk Island 
were recaptured and moved to the new park. The 
remainder of the 410 animals originally dehvered in 
1907, estimated to be only 48 head after the transfer 
in 1909 to Wainwright, formed the basis of the herd 
that still occupies part of Elk Island National Park. 
Later that year, Pablo delivered 218 head to 
Wainwright. In 1910, he was able to deliver 74 ani- 
mals and he made two shipments of seven bison each 
in 1912. In total, he was paid for 716 bison, 631 of 
which ended up in the Buffalo National Park. An 
additional 77 animals came from Banff National 
Park in 1909, and 30 came from the Conrad herd, 
which was also in Montana. 

According to one of the old files, the total number 
of animals introduced to the park was 738, but there 
is no record of the number of deaths or births during 
the years 1909 to 1911. Initially, the park, in total, 
occupied 159 square miles (a little more than 400 
square km). Later, more land was acquired on which 
to grow hay for winter feed. Experts in the Depart- 
ment of Agriculture estimated that its carrying 
capacity was about 7000 bison. 



3. Events at Buffalo National Park 

Rate of Growth in Numbers 

I have been unable to find a complete set of annu- 
al census Figures, either in the old files or other 
sources, but I found nine numbers for the years 1911 
to 1923 inclusive. (Table 1). Counts were apparently 
made in some, but not all years. 

Populations not under restraint tend to increase 
exponentially. A few hundred animals on a new 
range planned to support 7000 should be free of 
restraint. One of the properties of an exponential 
series is that the natural logarithms of the numbers 
lie on a straight line. The equation for the log^ trans- 
formed data is LN(Nt) = 4.562 + 0.192 t where 
LN(Nt) is the natural logarithm of the number of 
Bison "t" years after their introduction to the Park; 
4.562 is the logarithm of the number of Bison intro- 
duced to the park at time t = 0; and 0.192 (19.2%) is 
the annual rate of increase in numbers. The r^ for this 
equation is 0.995. 

Entrapment of other species 

When the initial perimeter fencing was completed, 
park personnel discovered that they had enclosed 
wild elk. Mule Deer, Odocoileus hemionus. Moose, 
Alces alces, and a few Pronghom Antelope, Antilo- 
capra americana (Lothian 1981: 29). I was suspi- 
cious about the antelope, but their presence was veri- 
fied by Rowan (1929: 359). Apart from competing 
with Bison for the available forage, there was a 
shght chance that the ungulates could have acquired 
diseases from contact with domestic cattle before 
they were trapped. In 1939, 1806 elk, 113 moose, 
and 242 deer that had been sharing the range with 
the Bison were slaughtered. Six per cent of the 
Moose and two individual deer "showed evidence of 
tuberculosis" (Lothian, 1981:37). 

Cross Breeding Experiments 

For several years The Department of Agriculture 
had followed the experiments of a Mr. Mossom 
Boyd in Ontario. Boyd had crossed Bison bulls with 
domestic cows and got fertile, hybrid offspring. The 
first generation hybrid offspring, when mated, also 
gave fertile offspring, which were named "Catalo" 
from the first three letters of "cattle" and last three 
letters of "buffalo." The name was later changed to 
"Cattalo." When Mr. Boyd died, the Department of 
Agriculture obtained his experimental animals and 
shipped them to an Agricultural Station at Scott, 
Saskatchewan. The agricultural scientists, anxious to 
do further experiments, entered into negotiations 



Table 1. Available Counts of Bison in Buffalo National Park, 1911 to 1923. 



Year 
Count 



1911 

73« 



1912 
994 



1913 
1188 



1914 

1453 



1915 
1640 



1916 

2077 



1919 
3830 



1921 
5052 



1923 
6780 



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143 



with the Dominion Parks Branch to obtain some 
Bison and space in Buffalo National Park so they 
could conduct more experiments in cross-breeding. 

At this point, Maxwell Graham, Director of 
Animals in the Dominion Parks Branch, enters the 
story. The salvaged files contain a memorandum 
from Graham to Mr. J. B. Harkin, Commissioner of 
National Parks, dated 27th June 1916. Graham 
reminded Harkin that they had discussed the ques- 
tion of cross-breeding "prior to my departure for the 
west . . . but the whole question was left in abeyance 
till Department of Agriculture was ready to take def- 
inite action." He went on to write: 

I was only made aware the other day that such action 
had been verbally undertaken during my last absence 
from Ottawa, and subsequent action in confirmation 
thereof through correspondence has only now been 
brought to my attention. 

I am only now aware that it is proposed that a portion 
of Buffalo Park, about six square miles, be fenced off for 
the purpose of keeping therein the Cattalo herd bought 
by the Department of Agriculture from the estate of the 
late Mr. Mossom Boyd; 

That the shipment of these animals took place on or 
about November 26th last from Bobcaygeon Ontario, to 
Scott Saskatchewan; 

That certain other bovines, being cows of the 
Galloway breed, are also to be imported, I believe from 
the east, and to be added to the Cattalo herd for experi- 
mental purposes. 

I now desire to point out that if such action is taken, 
the herd of bison, now over 2000 in number at Buffalo 
Park, will incur considerable additional risk of becoming 
infected with some variety of infectious disease. 
Graham admitted that there was already "a certain 
risk to be faced of some infectious disease obtaining 
a lodgement in our main herd" but he pointed out 
that most of the land around the park was under cul- 
tivation and such land carries on it "but very few cat- 
tle of any kind." He then expressed his fear that: 

We shall certainly run considerable risk, however, if 
we actually introduce into the park cattle in any quantity 
from Ontario. Some of these animals may, though them- 
selves immune, be carriers of disease from other herds. 

I would also point out that when an infectious disease 

is once brought into a large herd, the losses become very 

high, because it is difficult, if not impossible to check it 

after it has once obtained a foothold. 

He then went on to mention several such diseases 

— Hemorrhagic Septicemia. Anthrax, Foot and 

Mouth Disease — all of which had recently been 

discovered in the west. 

Finally, he most strongly recommended, in the 
same memorandum, that "some other area be found 
for the cross-breeding experiments... an area with 
natural drainage, and good running water and 
springs" instead of the "low land, stagnant water and 
liability to recurrent flooding of considerable areas" 
in Buffalo National Park. 

The Director of Agricultural Experimental Farms, 
J. H. Grisdale, lost no time in replying. His response 



is dated 5th July 1916 and required a little over three 
sheets of legal paper. His comments are in blue type, 
and the whole is peppered by comments in red type 
initialed M. G. (Maxwell Graham). The most impor- 
tant point comes near the end of Grisdale' s letter. 

I may conclude my remarks with respect to Mr. 
Graham's memorandum by saying that after having 
perused it most carefully, and after having discussed it 
with the Veterinary Director General, with the 
Pathologist of the Department of Agriculture, each of 
whom assures me that, with average precaution, the 
probabilities of any infectious diseases being transmitted 
to your buffalo through the proximity of the herd of cat- 
talo are practically negligible. 

Graham's response to that paragraph was directed 
to his own superiors. 

This, I consider completely absolves us from any pos- 
sible future criticism. Dr. Gordon Hewitt and myself 
each independently considered an added risk to be 
incurred & that is all I have contended, (signed) M.G. 
So the Bison, cattle, and cattalo of the Agriculture 
Department were established on 6 square miles of 
park land. Only a single fence, which permitted nose 
to nose contact, separated them from the Bison herd. 
Later, they were joined by several Yaks (Bos miitus). 
Grisdale guaranteed that all animals would be tested 
for diseases before being released inside the cattalo 
pen. Did any infectious diseases come with the 
experimental animals? We will never know for sure, 
but Graham was to raise the question again several 
years later. 

The cattalo episode did raise other questions. In a 
six-page (legal size) memo dated 7th July 1916 
Graham reviewed the precautions already taken 
since 1912 and what was still needed. He made it 
plain that a major requirement was some means of 
segregating individuals by means of a system of cor- 
rals and cross fences. He had first made that request 
in 1914, but his request was turned down in a letter 
from Head Office to the Superintendent of Buffalo 
Park, dated 19th December 1914. The letter stated 
simply that "[T]he expense of constructing further 
fenced subdivisions would appear to be out of the 
question at present." He fared no better in 1916. 

First Indications of disease 

The threat of disease was always a major concern 
for Graham because of his position as Chief of the 
Animal Division. His immediate superior, to whom 
he reported, was J. B. Harkin. Commissioner of 
Dominion Parks. There are a number of lengthy 
memoranda among the files from Graham to Harkin, 
urging that action be taken to reduce the risk of a 
serious disease outbreak. Virtually all of his plead- 
ing, such as the need for a system of corrals cited 
above, fell on deaf ears. 

h was almost 1917 before the first case of tuber- 
culosis was discovered. On 20 December 1916 a 
post mortem inspection of a dead bull revealed 



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lesions typical of tuberculosis. In January 1917 a 
visual inspection of the herd was ordered. Help was 
received from an inspector from The Agriculture 
Department. Out of the herd of 2 000+, only two or 
three animals were judged to be in poor condition. 
They were segregated from the main herd, but appar- 
ently not tested or slaughtered and examined post 
mortem. 

Veterinary Inspector Gillam wrote to Graham and 
recommended an enclosure with corral and a 
squeeze chute, "which would prove useful should it 
at any time be deemed advisable to vaccinate against 
Blackleg, Anthrax, or Contagious Abortion." (The 
last of these is also known as Brucellosis.) No corral, 
let alone a squeeze chute, was forthcoming. 

No confirmed new case of tuberculosis showed up 
until 20 March 1919 when a dead cow was found, 
examined by the local Veterinarian, and diagnosed 
as having died from tuberculosis. The cow was 
thought to be about 15 years old. 

On 22 April 1919, a 4-year-old bull was killed and 
a post mortem examination revealed "traces of T.B." 

On 13 April 1920, "an apparently healthy buffalo 
bull selected for slaughter for the Hudson's Bay 
Company banquet, was found to be tubercular." 

In spite of the evidence cited above, the Deputy 
Minister of the Department of Interior denied, in an 
interview published by the Montreal Gazette for 2 
December 1922, that there was any tuberculosis in 
Buffalo National Park. 

Incontrovertible Evidence of Disease 

The first slaughter of Bison at Buffalo Park took 
place in February 1923, when 264 Bison were killed, 
butchered, and examined by a dominion veterinary 
inspector. Dr. A. E. Cameron (later the Veterinary 
Inspector General) and Dr. Seymour Hadwen, then 
of the Ontario Research Foundation, were also pre- 
sent. Cameron (1923) published a small paper enti- 
tled "Notes on Buffalo: Anatomy, Pathological 
Conditions, and Parasites" without even mentioning 
tuberculosis. Hadwen withheld publication of his 
observations until 1942. 

No details about disease are in the files that I 
have, but Lothian (1981: 32) wrote that "The inspec- 
tors found that 75 percent of the animals slaughtered 
(in 1923) had some form of tuberculosis lesion." He 
did not mention, though, that the sample was strong- 
ly biased toward old males and thus was probably 
not representative of the herd as a whole. 

Lothian goes on to say (Page 32): "Dr. Hadwen 
recommended the elimination of the herd to avoid 
the spread of infection, but the proposal was not 
acceptable to park authorities." The following win- 
ter, 1923-24, 1847 bison were killed, but there is no 
record in the files of the occurrence of any disease. 
Dr. A. E. Cameron (1924) published a second paper 
entitled "Some Further Notes on Buffalo" In which 



he disposed of tuberculosis in a single sentence on 
the last page: "Tuberculosis has been found in buffa- 
lo, as is common in wild animals in captivity." 

Lothian's only comment was (page 32): "The 
presence of disease was withheld from public knowl- 
edge as a matter of departmental policy." Apparently 
it was still a matter of departmental policy in 1981 
when Lothian's book was published. 

Instead of informing the public about the health of 
the herd, the government distributed pamphlets con- 
taining recipes for preparation and cooking of buffa- 
lo meat, whether from healthy or diseased animals. 

When Graham, who had transferred from the Parks 
Branch to the Northwest Territories and Yukon 
Branch at the beginning of 1922, learned the results of 
the slaughter he wrote a memorandum dated 13 April 
1923 to W.W. Corry, Deputy Minister, Department of 
the Interior. It was marked "Personal and Con- 
fidential." The pertinent points raised by Graham, with 
my comments in square brackets, were the following: 
As the official particularly charged with the adminis- 
tration of the buffalo in National Parks from January 
1912 to January 1st, 1922, I am naturally concerned to 
hear that as a result of investigations made by Dr. 
Seymour Hadwen, he has reported the herd at Buffalo 
Park as being seriously infected with Tuberculosis 

I am fully aware that under the circumstances, once 
Dr. Hadwen' s report is made public, I, as former 
Director of Park Animals, must expect considerable crit- 
icism. Knowing as I do, and as our records will plainly 
show, that every effort to prevent such a calamity has 
consistently and all along been made by me, I personally 
do not fear such criticism. 

But the loss of your confidence in me I should feel 
very keenly, and in order to forestall such I wish to bring 
to your attention the following facts. 

1st. During my tenure of office, so far as I am aware, 
there have only been four cases of tuberculosis, definite- 
ly so pronounced, in Buffalo Park. [The four cases 
already noted above.] 

2nd. During the slaughter of surplus buffalo in 
September 1922, Doctor Hadwen I understand, reports 
lesions in organs examined from the condition of which 
he estimated that not more than six years could have 
elapsed since infection took place. And, so I understand, 
infection in most cases was of a more recent date. 

3rd. Our records will show that as far back as 1914, I 
strongly recommended that the Buffalo in Buffalo Park 
should be allowed to freely range during winter, instead 
of being confined in "winter quarters", and I stated "The 
danger of infectious or contagious diseases attacking the 
herd would not be as great (if allowed to winter in the 
main park) as if the herd were all kept together in con- 
fined quarters" (or confined feeding area.) 

My recommendation was turned down, without being 
referred to you, and it was not till July 1917 on the 
advice of Doctor Hargrave. Chief Inspector for Alberta, 
that action was taken to carry out my recommendation 
of 1914. 

I would suggest that Doctor Hadwen, as an unbiased 
authority, might be asked his opinion on the points 



I 



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Fuller: Canada and the Buffalo 



145 



raised in my memorandum of June 27th, 1916, and 
whether or not in his opinion the reconrunendations made 
in my memorandum re tuberculosis of March 19th, 1919 
were sound. 

In any case, these latter were not acted on except in so 
far as reference is made to them in the attached memo- 
randum of April 1st, 1919 by Doctor Torrance. 
The part of Dr. Torrance's letter to which he 
refers seems to be the following: 

The whole question is one of providing plenty of 
nourishment and fresh air. The buffalo, I understand, are 
constantly out in the open, and they should always be 
provided with an abundance of nutritive food so that 
they may be maintained in as vigorous a condition as 
possible. 

The next, and last, point raised by Graham must 
be read in association with his 2nd point. 

It only remains to be added that my attempt in 1916 
to prevent what I then considered, and still consider, an 
uncalled for additional risk to the buffalo. . .was met with 
by unmerited censure and ridicule. 

When in September 1922 Doctor Hadwen made his 
investigation of the herd in Buffalo Park, just six years 
had elapsed since, against my protest, the cattalo and 
domestic bovines were first placed in Buffalo Park. 
Graham is clearly making a connection between 
arrival of the Bison, cattle, and yaks in 1916 and Dr. 
Hadwen' s finding that none of the cases of tubercu- 
losis were older than 6 years, i.e. acquired no earlier 
than 1916. He does not go quite so far as to make an 
outright accusation. 

Without, however, actually blaming the cattalo and 
domestic bovines in the experimental herd for the seri- 
ous condition of our buffalo in Buffalo Park, it would 
seem that my modest expression of opinion as to the 
added risk incurred by insistence in selecting the place 
for cattalo experiments where it is was fully justified. 
Written in pencil across the upper left hand comer 
of the first page of the memorandum is the following: 
"Mr. Graham, I don't see how you can be blamed 
in this matter. W.W.C." The initials are those of 
W. W. Cory, Esq., C.M.G., Deputy Minister, 
Department of the Interior, to whom the memoran- 
dum was addressed. 

There is also a short memorandum directed to Mr. 
Graham and signed by O. S. Finnic, Commissioner 
of the Northwest Territories and Yukon Branch, 
Department of the Interior, to which Graham now 
belonged. It read: 

I read over, with much interest, the correspondence 
hereunder. The evidence shows conclusively that you 
always entertained apprehensions as to the safety of the 
herd at Wainwright — even in the face of expressions to 
the contrary from those in high authority. They cannot 
lay at your door the charge that you failed to advise 
them of the dangers of disease. 
I interpret this to be, on the part of the government 
at least, a clear exoneration of Maxwell Graham. 

The Horns of a Dilemma 

Even before 1920, the relentless pressure of expo- 
nential increase in the size of the herd, coupled with 



deterioration of the range, created a serious problem 
for Buffalo Park managers on the site, senior civil 
servants in Ottawa, and ultimately, elected politi- 
cians. Something had to be done — but what? The 
only options appeared to be to continue with annual 
slaughters, or slaughter of the entire herd, or transfer 
the surplus to some other suitable location. 

Hewitt (1921: 136), whose book was published 
posthumously, had some further suggestions. One 
was to "establish small parks in other parts of the 
Prairie Provinces, where small herds could be main- 
tained, which would make the bison accessible to 
people". He suggested also (136) "[E]very large city 
should have its zoological park." And finally he set 
out his view on the possibihty of domestication. 

"The greatest value of the buffalo, however, lies 
in the possibility of its domestication... the greatest 
need of the prairie provinces is an increase in its 
beef-producing capacity. The buffalo is an animal 
which offers great possibilities, being pre-eminently 
suited to prairie conditions, and at the same time it 
produces a robe of no small commercial value" 
(136). At the time when this was written, no one 
seemed to take domestication seriously, and disease 
did not seem to be a barrier. Today, in North 
America, there are many more Bison on private 
ranches than in parks and reserves. 

When the slaughter of nearly 2000 bison in 1923 
was made public there was a strong reaction against 
further slaughters. Given the strong public objection 
to a simple herd reduction, the Parks Branch felt that 
total elimination of the Buffalo National Park herd, 
as suggested by Hadwen, was out of the question. 
Dr. C. H. D. Clarke, who salvaged the files, was also 
in favour of total slaughter and restocking with 
healthy Bison from Elk Island National Park. 

Wood Bison 

In January 1772 Samuel Hearne and his Indian 
companions crossed Great Slave Lake from north to 
south. Hearne (1795: 250) wrote: 'immediately on 
our arrival on the south side of the Athapuscow [now 
Great Slave] Lake, the scene was agreeably altered, 
from an entire jumble of rocks and hills, for such is 
all the land on the North side, to a fine level country, 
in which there was not a hill to be seen, or a stone to 
be found." As they made their way up the lowlands 
on the eastern side of the Slave River, he noted that 
"Buffalo, moose, and beaver were very plentiful...." 
Hearne (1795: 251) also noted that "The buffalo in 
those parts, I think, are in general much larger than 
the English black cattle: particularly the bulls, 
which, though they may not in reality be taller than 
the largest size of the English oxen, yet to me always 
appeared to be much larger. In fact, they are so 
heavy, that when six or eight Indians are in company 
at the skinning of a large bull, they nc\cr atlcmpl to 
turn it over while entire, but when the upper side is 



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The Canadian Field-Naturalist. 



Vol. 116 



skinned, they cut off the leg and shoulder, rip up the 
belly, take out all the intestines, cut off the head, and 
make it as light as possible, before they turn it to 

skin the under side The head of an old bull is of 

great size and weight indeed: some which I have 
seen were so large, that I could not without difficulty 
lift them from the ground." Heame was thus the first 
European to see and describe the wood buffalo. 

When I was at Fort Smith in the 1940s and 1950s 
I took part in the skinning of many a Bison, and I 
watched many Indians and Metis working in pairs. I 
never saw them lighten the load before turning over 
a Bison in Wood Buffalo National Park. 

Heame was not usually given to exaggeration, as 
were many explorers of his day, so I take his obser- 
vations as evidence that the Wood Bison were con- 
siderably larger than those of the plains. 

Other early explorers saw Wood Buffalo in the 
course of their journeys. While visiting Forts 
Chipewyan and Wedderburn, on Lake Athabasca, 
Captain John Franklin (1823:154) noted that "The 
traders here also get supplied by the hunters with 
buffalo and moose deer meat (which animals are 
found at some distance from the forts). . ." 

After descending the upper Slave River, and nego- 
tiating the rapids above the present Fort Smith, 
Franklin ascended the Salt River as far as the salt 
springs, where his men collected salt for the rest of 
their journey. On leaving the springs (Page 107) they 
"perceived a buffalo plunge into the river before us." 
The unfortunate animal was immediately shot and 
kept the entire crew in meat until they reached Fort 
Resolution on Great Slave Lake. While there he was 
informed (199) that the residents "Procure moose, 
buffalo, and rein-deer meat occasionally from their 
hunters; but these animals are generally found at the 
distance of several days' walk from the forts." 

Joseph Sabine, a zoologist of Franklin's time, pre- 
pared The Zoological Appendix (Number V), to 
Franklin's narrative. About the Bos Americanus or 
American Buffalo, he wrote (1923: 688) "... they are 
extremely numerous on the plains of 
Sashatchawan,(sic) and are also found, though less 
plentifully, in the woods as far north as Great Slave 
Lake; a few frequent Slave Point, on the north side 
of the lake, but this is the most northern situation in 
which they were observed by Captain Franklin's 
party." 

The reference to Slave Point, on the north side of 
the lake, has been cited by several other authors, 
who, apparently, did not verify its location. Slave 
Point was said to be composed of limestone. The 
only Slave Point on modern maps is at approximate- 
ly 61° 10' N and 1 15° 55' W on what 1 would call the 
west shore rather than the north shore. No Slave 
Point appears on Franklin's map on the true north 
shore which, in any case is composed of 
Precambrian granites, not flat limestone. 



In the middle of February 1 890, Warburton Pike 
(1892: 143) set out from Fort Resolution on a buffa- 
lo hunt with an Indian guide named Francois. His 
assessment of the distribution of bison was as fol- 
lows: 

"Scattered over this huge extent of country are 
still a few bands of buffalo. Sometimes they are 
heard of at Forts Smith and Vermillion, sometimes at 
Fort St. John close up to the big mountains on Peace 
River, and occasionally at Fort Nelson on the north 
branch of the Liard." 

After reaching the end of Francois' road, and 
walking until noon, they came on a band of eight 
bison and killed one cow. A few weeks later he 
made a second hunt, again with Francois as his 
guide, but they saw no trace of bison. Pike (1892: 
43) appears to have been the first to attribute the size 
of the wood buffalo to environmental factors as the 
following quotation shows: 

"These animals go by the name of wood buffalo 
and most people are of opinion that they are a dis- 
tinct race from the old prairie buffalo so numerous in 
bygone days; but I am inclined to think that the very 
slight difference in appearance is easily accounted 
for by climatic influences, variety of food, and the 
better shelter of the woods."" The italicized words are 
one of the untested hypotheses that will play an 
important role in what happened later. 

In 1892, Frank Russell, a young academic at the 
University of Iowa, began his journey into the far 
north. On 9 January 1893, Russell set out from Fort 
Resolution, guided by Fran9ois who had guided 
Pike. As Pike had done, he began his journey on the 
Little Buffalo River and later turned off to the south- 
west. On the fourth day they reached what he called 
(Russell, 1898: 103). "the northern limit of the buf- 
falo range, perhaps fifty miles south of the Great 
Slave Lake." They saw no buffaloes and had to 
return empty handed after 13 days on the trail. 

Early in the spring of 1902, Charles Camsell was 
asked to search the country south and west of Fort 
Smith for Wood Bison. At that time he was a very 
junior member of the Geological Survey, which was 
in the Department of the Interior. Before he retired, 
at age 70, he had risen to Deputy Minister of the 
Department of Mines. His journey is set out in his 
memoirs, Camsell (1954). He and his assistant 
launched their canoe at Athabaska Landing (now 
Athabasca) and paddled down to Smith Landing 
(now Fort Fitzgerald). From there they went by ox 
cart to Fort Smith. From Fort Smith, they made two 
short sorties and saw only one small bison herd and 
many tracks. Their final trip took them to the Peace 
River by way of the Salt River, the upper Little 
Buffalo, and the Jackfish River. They began that 
journey on 2 August and reached the Peace a whole 
month later. Camsell estimated that the number of 
Wood Bison was probably about 300, but he gave no 
data of any kind to support his estimate. 



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147 



Ernest Thompson Seton and Edward A. Preble set 
out for the north, by canoe, in 1907. Seton could not 
have chosen a better companion than Preble who, 
with his brother, covered about 1200 miles by canoe 
in the Hudson Bay region in 1900, and in 1903-04 
travelled several hundred more miles in the 
Athabaska-Mackenzie Region. 

En route to the tundra, Seton and Preble paused at 
Fort Smith for a buffalo hunt. Their guide led them 
to a group of 13 bison the first day (page 48) and 22 
the second day (page 52). An account of Seton' s trip 
was pubUshed in 1911 (Seton 1911) and repubhshed 
in 1943. 

It is apparent from this brief survey that Bison 
hved at least as far north as the south shore of Great 
Slave Lake and perhaps further. It is also apparent 
that by the end of the 19th century, their numbers 
were rather small, and the largest herd was thought 
to live in the angle formed by the junction of the 
Peace and Slave Rivers. 

Maxwell Graham visited the area south of Fort 
Smith in 1912. On his return to Ottawa he wrote a 
memorandum, dated 16th October 1912 to J. B. 
Harkin. He dealt briefly with the character of the 
country through which bison ranged, the length of 
time that they are known to have lived in the region, 
and the size and type of "these pure-blood bison." 
He argued that although ample range is critical for 
their survival, ample range alone is not enough. 
They also need other kinds of protection. Graham 
suggested that protection would be easier and less 
expensive if the various small groups of bison could 
be herded into a single reserve. He felt that the ideal 
location for such a reserve was the junction of the 
Peace and Slave Rivers (Figure 1). He noted that: 

A line drawn eastwards from Peace Pt. upon the 

Peace River, would intersect the Great Slave River about 

20 miles below the junction with the Peace. If a suitable 

fence were constructed along this line, a distance of 

25-30 miles, the Buffaloes could gradually be worked 

from the North into this peninsula where there would be 

ample range for a number of years. 

Graham thought that "herders" could turn back 

any Bison that sought to cross the Slave or the Peace 

River in winter over the ice, but he neglected the 

possibility of escape by swimming across the rivers 

in summer, which Bison do with ease. 

Graham also quoted from what he called the 
"Smithsonian report." (See Homaday 1889). 

'There is reason to fear that unless the government 
takes the matter in hand and makes a special effort to 
prevent it, the pure-blood bison will be lost irretrievably 
through mixture with domestic breeds, and through in- 
breeding; this latter causes eventually, loss in size and 
final sterility.... As I have already stated no suitable 
stock can now be procured from any of the herds in the 
United States, a large number of these are not pure- 
blood, some which are, have deteriorated in size owing 
to too close confinement and in-breeding." 



That most of the herds in the United States are not 
pure-blood has recently been confirmed by mito- 
chondrial DNA sequencing (Polziehn, et al., 1995.) 

Graham's memorandum closes with the following 
paragraph. 

Such being the case I again urge that the future 
administration of this Northern Canadian Herd, the last 
and best of the buffalo, be entrusted to this Dominion 
Parks Branch, and that provision be made in the esti- 
mates to provide for the enclosure proposed above, a 
plan of which is appended herewith and that a vigorous 
policy as to this herd's protection be instituted, and a 
capable superintendent be appointed who shall have the 
interests of this Branch at heart, and who would also 
understand and appreciate the policy of the Dominion 
Parks Branch." 

No action was taken on Graham's memorandum. 
The need for a reserve of some kind was raised again 
in 1914 at a meeting of the Council of the Northwest 
Territories, which met in Ottawa in those days, but 
again it was not acted upon. 

Camsell made three more short trips into the 
Wood Bison range during the summer of 1916 at the 
request of the Parks Branch of the Interior Depart- 
ment. A type-written copy of his report is among the 
salvaged files. On page 1 Camsell wrote: 

The wood bison of northern Alberta and the adjacent 
portion of the Northwest Territories are in nvo separate 
bands occupying two distinct ranges, and there does not 
seem to be at present or within recent years any migra- 
tion of buffalo from one range to the other. ... A belt of 
muskeg country 30 to 40 miles wide separates the north- 
em from the southern range and prevents migration from 
one herd to the other except by way of the Salt plain. 
The italicized portion is the second untested 
hypothesis that was to play a role later on. 

The second important conclusion in Camsell' s 
report is that most of the Wood Bison were in an 
area extending from the Peace River north to latitude 
65° (clearly a typographical error) between longi- 
tudes 112° and 1 13°. On page 5 he suggests a game 
reserve bounded by 60° on the north, 1 13° 20' on the 
west, latitude 59° 10' on the south, and Slave River 
on the east. No action was taken on his recommen- 
dation for a reserve. In 1918, another recommenda- 
tion to create a park was also turned down. 

In 1922, Maxwell Graham revisited the range of 
the Wood Bison. His report, entitled "Some obser- 
vations on the wild buffalo in Northern Alberta and 
the Fort Smith District, Northwest Territories," fills 
5 sheets of legal paper and is dated 23 November 
1922. The report begins with a notation that "true 
buffalo" are found only in Africa and Asia, and that 
the "buffaloes" of Europe and North America are 
properly known as "bison." There is only one 
species of Bison in Europe. Bison bonasus. There is 
also only one species in North America, Bison 
bison, which was thought by many to have two 
subspecies. 



148 



The Canadian Field-Naturalist 



Vol. 116 



Graham wrote: 

Both the writer and his colleague are in a position to 
state that our Northern wood-buffalo are larger, darker, 
and handsomer specimens than those which formerly 
ranged the plains far to the south. 
He goes on to say, however, that: 

in the writers (sic) opinion, whatever differences 
there are between it and those of the plains, is (sic) 
entirely owing to environment. 

Graham clearly agreed with Pike's untested 
hypothesis. It is not clear whether or not Graham 
believed that Wood Bison would revert to Plains 
Bison if they were moved to central Alberta, or con- 
versely, Plains Bison would come to resemble Wood 
Bison once they were moved to the wood bison envi- 
ronment. On page 2 of the memorandum Graham 
wrote: 

The buffalo at Elk Island Park are supposed, in the 
aggregate, to be the best in any park, nevertheless, with- 
out meaning to disparage this fine herd, the writer and 
others who were with him on the wood-buffalo range 
must state that in size, thriftiness, stamina and general 
appearance the wild wood-buffalo is in a class by itself 
and quite unapproachable. 

The use of "in a class by itself suggests that they 
were worth preserving whether or not they were for- 
mally recognised as a subspecies. 

Graham made his headquarters at Pine Lake in 
1922. A wagon road connected Ft. Smith and Pine 
Lake. About two miles before it reached the lake, the 
trail crossed the upper Salt River. When I worked in 
WBNP in the late 1940s, the old trail and the cross- 
ing were still visible and I visited the old crossing, 
and saw the tree in which Graham had a platform 
from which to watch Bison. One of the first "buffalo 
rangers," Billy McNeil, was my informant. 

Here are some excerpts from notes Graham made 
on August 12, 1922. 

The most successful day yet. Dull and raining in the 
morning, but clearing towards noon. Stewart and I start- 
ed (on foot) for Salt River Crossing. At Wallows on the 
way I spotted a huge bull lying down, however, he wind- 
ed us and got away before we could get a picture. On 
arrival at Salt River Crossing, Stewart climbed a leaning 
tree and I posted myself across the river under a small 
tamarac,(sic) in full view of open muskeg. We saw and 
got pictures first of a herd of 1 6 buffalo. In about half an 
hour another herd of about the same size came out of the 
forest, of which we also got some pictures. Stewart fol- 
lowed these in hopes of geUing some close-ups, none of 
them having seen or scented us... Shortly after, Stewart 
returned not being able to catch up the buffalo in the 
bush beyond river (sic), and it was then that I heard 
more coming... So far as I could estimate there were 9 
in this herd, one immense bull, two cows, two 2 year 
olds, 3 yearlings and I very small calf, whose skin was 
throughout of a faint yellowish white colour. 
All in all, Graham and his assistant saw and 
counted 1 14 bison, which was more than all other 
visitors mentioned above had seen. He also ventured 
to estimate the total number. His estimate was based 
upon 



...the finding of old trails and old wallows being 
reopened as well as the presence of new ones, and from 
the number of young stock seen it is safe to state that the 
wood-buffalo are increasing, and from all the evidence 
our party was able to investigate, at a conservative esti- 
mate, the number of buffalo in the southern range may 
be put down as 1000. 

Graham also argued again for more protection 
from poachers, forest fires, and Wolves (Canis 
lupus), in the form of an enlarged corps of rangers 
(now wardens). The last paragraph in the report sug- 
gests that Graham had some inside information. 

Both for the protection of the buffalo as well as valu- 
able forms of other wild life, it is expected that steps will 
shortly be taken to conserve for posterity the range on 
which the buffalo roams. 

Another memorandum dated 13th November 1922 
is entitled "Reasons Why the Wood Buffalo of 
Northern Alberta and Northwest Territories should 
be preserved." 

These buffalo are the last of their species living to- 
day under absolutely free and wild conditions. 

They are the finest specimens of their species, superi- 
or in pelage, size, and vigour to those of the plains, 
whose descendants to-day exist in our parks. 

Owing to certain physiological causes buffalo in cap- 
tivity deteriorate in size and stamina, further, when in 
confinement their natural increase contains an undue 
proportion of males to females, which unless steps are 
taken to remedy this surplusage (sic) of males is apt to 
bring about tuberculosis and kindred diseases. 
We know now that sex is determined by presence 
or absence of a Y chromosome, and it is not clear 
how the sex ratio is involved in the spread of dis- 
ease. 

The time is approaching when an infusion of new, 
unrelated, blood will be needed by our herds in the 
National parks, and it is only from the northern herd that 
such infusion can be obtained. 

Looking to the future success of the experimental 

cross-breeding between buffalo and domestic bovines, it 

is imperative that a reserve stock of pure blood bison of 

the highest potency should be kept in reserve, so that the 

ultimate fixed type of new range animal may continue to 

pass on to successive generations the prepotent qualities 

of the true bison, viz hardiness, thriftiness, a valuable 

robe and first class beef qualities. 

A second typescript is also among the files. It may 

be a draft of "Canada's Wild Buffalo," which was 

published by the Department of Interior in 1923. It 

contains more information than the memorandum 

about the nature of the country, such as meadows, 

muskegs, sloughs, poplar ridges with edible grasses, 

jack pine ridges that have a scanty ground flora, and 

sink-holes that develop in the gypsum that underlies 

much of the area. 

Associated with the report and manuscript was a 
letter, dated 22 November 1922, to Graham from 
R. M. Anderson, who was then Chief, Division of 
Biology, Victoria Memorial Museum. His letter 
reads, in part, as follows. 



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149 




3 



♦^ #% «i.i^j»»i» 




Figure \. Maxwell Graham's original map showing the area he proposed as a reserve for wood buffalo in the 
angle between the Peace River, which comes in from the southwest, and the Slave River which exists to 
the north. The line near the top is the 6()th parallel. The grey patches in the lower right arc water bodies. 



I have carefully read over the manuscript of your 
description of the wood buffalo range, observations on 
the habits of the wood buffalo, and reasons why the 
wood buffalo of northern Alberta and Northwest 
Territories should be protected. It is very interesting as a 



very up-to-date summary of the wood buffalo situation 
as well as a comparison of the wood buffalo with the 
more southern herds, and there is really nothing that I 
can critici/e in the paper. 

The desirability of keeping a reserve stock oi pure 



i 



150 



The Canadian Field-Naturalist 



Vol. 116 



blood buffalo, living under free, natural conditions, is so 
obvious as to need no argument.... 

Thanking you for your kindness in loaning me the 
report, I remain 

Yours sincerely, 

(signed) R. M. Anderson 

About one month after Graham submitted his 
reports, the steps that he had predicted came to pass. 
An Order in Council established Wood Buffalo 
National Park (WBNP) in December 1922. The stage 
was now set for the two streams we have been follow- 
ing to come together. Suddenly, 10 500 square miles 
of sparsely populated wood bison range became avail- 
able to receive the excess in the National Park at 
Wainwright. 

The Transfer Goes Ahead 

The following information is taken from a docu- 
ment assembled by Col. J. P. Richards dated 6th 
May 1923. Col. Richards held a senior position in 
the Department when I joined it in 1947, and I found 
him to be an honest, dedicated public servant. The 
title of the document is "Summary of correspon- 
dence dealing with the transfer of the buffalo from 
Wainwright to Wood Buffalo Park." Notations in 
square brackets are my conmients. 

1. The decision makers 

Under date of 23rd May, 1923, the Commissioner of 
Canadian National Parks advised the Department that 
there were too many buffalo in Wainwright Park and 
some arrangement would have to be made to reduce the 
number. 

The Department suggested that instead of slaughter- 
ing excess buffalo it might be desirable to transfer some 
of the healthy [?] young stock to the Wood Buffalo Park 
and it was arranged to call a conference to discuss the 
proposal. The conference was held on the 30th May and 
the following extracts which are taken from a memoran- 
dum dated 5th June, 1923, outline the decisions arrived 
at. 

Under date of May 30th. 1923, on Wednesday at 3 

p.m. a conference was held at the Commissioner's 

office, at which were present Mr. W. W. Cory, acting in 

a dual capacity as Deputy Minister of the Department 

and Commissioner of the Northwest Territories, Mr. 

O. S. Finnie, Director of the Northwest Territories, Mr. 

J. B. Harkin, Commisssioner of National Parks, Doctor 

F. Torrance, Veterinary Director General, Department of 

Agriculture, and Mr. A. Smith, Superintendent of the 

Wainwright Park. 

It is important to note that Maxwell Graham's name is 

absent from the list of participants. He took no part in that 

conference. 

The question before the Conference was concerned 
with the disposal of buffalo surplus to the capacity of the 
Wainwright Park, and also the advisability of introduc- 
ing these surplus buffalo into the Wood Buffalo Reserve. 
The question was rendered more complex owing to 
the fact that tuberculosis had manifested itself among the 
buffalo in the Wainwright Park. [My emphasis] 



The conference was opened by the Deputy Minister' s 
statement that Mr. Harkin had informed him something 
must be done to relieve the growing congestion of buffa- 
lo at the Wainwright Park, that the slaughter of a large 
number of buffalo had been suggested, but he disliked 
the idea of such slaughter and had thought a way out of 
the difficulty could be found by transplanting the surplus 
Wainwright buffalo into a new environment away from 
all danger of communicable disease. [Except what they 
took with them, of course.] 

Such an area was to be found in that area selected by 
the buffalo themselves, whose descendants are now 
known as wood buffalo and whose habitat near Fort 
Smith has recently been created a park. Addressing in 
particular Doctor F. Torrance, the Deputy Minister 
requested his opinion with respect to the chances of 
recovery from tuberculosis if buffalo infected with this 
disease were transplanted in the Fort Smith district, and 
if in his opinion there would be any serious menace to 
the wood buffalo through such action being taken. 

Doctor Torrance was of the opinion that although in 
some less advanced cases an improvement might be 
looked for, the transplantation of any considerable num- 
ber of the infected herd into the range of one, presum- 
ably, not infected would be extremely hazardous. 

It was then finally decided that selected young stock, 
not over one year old, should as soon as possible, be seg- 
regated, corralled, and squeeze-chuted, in order that each 
one might be subjected to the intradermal test. Those 
passing the test to be shipped by rail to Peace River 
Crossing, thence by scows, through the [Vermilion] 
chutes, to Peace Point and liberated in the Wood Buffalo 
Reserve. [Peace Point is an outcrop of limestone at least 
15 metres high!] 

Dr. F. Torrance of the Department of Agriculture, 
who was present at the conference, expressed his 
opinion about the disease problem in a letter to the 
Department on the 30th of the same month. 

As regards Tuberculosis, the suggestion has been 
made that some of the younger buffalo should be trans- 
ported to the range of the wood buffalo in the north and 
permitted to mingle with them. This proposition is 
objectionable from a health point of view, in that it 
would be almost certain to carry infection to this herd of 
wood buffalo, which presumably is at present free from 
this disease. [My emphasis] 

If this proposition were, however, modified and prep- 
aration made so that young animals up to the age of 
yearlings only were transferred, and that these animals 
were, previous to transference, submitted to the tuber- 
culin test, so as to eliminate any that reacted, much of 
the objection would be removed. 
Note that Torrance did not give his outright bless- 
ing to use of the tuberculin test. As late as the 1950s 
at least, the tuberculin test gave some false negatives 
so Dr. Torrance would have known that a single test 
would not have detected all reactors. 

Under date of the 13th November, 1923, the 
Edmonton Journal published a despatch from Ottawa 
in which it was stated that the Minister of the Interior 
had announced that it was the intention to transport 
two thousand buffalo calves from Wainwright to 
Wood Buffalo Park. 



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A second Parks Branch Memorandum dated 20th 
November, 1923, dealt only with the steps to be 
taken to prevent injury to the animals in transit from 
Wainwright to WBNP. The Commissioner, 
Canadian National Parks Branch, on the 22nd of the 
month, commented upon the memorandum referred 
to in part as follows: 

There is one subject which has been discussed in the 
department which is not mentioned in Mr. Nagle's mem- 
orandum, that is the possibility of the transfer of 
Wainwright buffalo to the wood buffalo range resulting 
in the introduction of tuberculosis in the wild herd. You 
will remember that last spring we had a conference with 
Dr. Torrance upon this subject and that the Doctor 
would not commit himself concerning this aspect of the 
situation. 

A letter dated 27 November 1923 from J. B. 
Harkin to Edmund Seymour, President of the 
American Bison Society was pubhshed in the Report 
of the American Bison Society for 1922-23 page 32. 
It read as follows: 

"Since last winter the Department has had under con- 
sideration a proposal to make shipments of buffalo from 
the surplus of the Wainwright herd to Fort Smith coun- 
try, now occupied by the so-called wood bison. It is the 
intention of the department to make some experimental 
shipments next Spring but I doubt very much whether 
the number will be as large as one thousand. Difficulties 
in connection with the transportation of adult buffalo to 
such a remote area are so great that it is improbable that 
any attempt will be made to send adults to the north. 
Yearlings appear to be the only type that could be han- 
dled on such a project. Just what number will be sent has 
not been decided. I do not think a decision can be 
reached until next spring. The only established fact in 
connection with this subject is that the Department's 
present intention is to make an experimental shipment." 
Nowhere does the summary of discussions on 
movement of Bison mention an "experimental ship- 
ment." So why did Harkin write the letter? It could 
have been to alert Seymour with the hope that he 
would protest the shipment in the name of the Bison 
Society. It is even possible that Maxwell Graham 
drafted the letter for Harkin 's signature. The copy of 
the Bison Report for 1922-23 that arrived with the 
files has Graham's name and departmental address 
on the cover. 

There followed a copy of a report by an Inspector 
Waddy on the prevalence of tuberculosis in 46 bison. 
According to the file that lists all slaughters from 
1923 to 1937 there was "no slaughter" in 1924-25, 
but a copy of the report, dated 10th Jan'y 1924, was 
sent to the Veterinary Director General, and a copy 
is among the salvaged files. The report begins with 
an arbitrary, but adequate, definition of three age 
classes. The definitions are as follows: 

With not more than one pair of permanent teeth up — 

under 5 years 

With all permanent teeth up but not worn at the edge — 

5-10 years 

With all teeth in wear — over 10 years. 



Table 2. Prevalence of tuberculosis in a sample of 46 bison 
examined by Dr. Waddy in January, 1924. 



Age 


Number 


Number 


Percentage 


Class 


Slaughtered 


Positive 


Positive 


1 


15 


2 


13.3 


2 


13 


9 


69.2 


3 


18 


16 


88.9 


TOTAL 


46 


27 


58.7 



Table 2 shows Dr. Waddy' s findings. 
Waddy remarked "I think this plainly shows the 
prevalency of generalized T.B. in the old animals." 

The numbers certainly speak for themselves. 
Prevalence increases with the age of the animal, and 
more than half of the animals in the sample were 
infected with tuberculosis. The findings also help to 
explain the high incidence of tuberculosis in the 
1923 slaughter that was biased toward old males. 
The Richards memorandum went on as follows: 

This report was received in the Department at the 
time consideration was being given to the proposal to 
transfer a number of Wainwright Buffalo to Wood 
Buffalo Park. It will be observed that fifteen animals five 
years of age and younger were examined by Inspector 
Waddy and only two showed slight evidence of tubercu- 
losis. No animals over the age of three years were trans- 
ferred to Wood Buffalo Park and the number of three- 
year olds that accompanied the shipment was less than 
five per cent of the total. 

At a conference held in the Deputy Minister's office 
on the 9th April, 1924. it was decided to transport 500 
yearling buffalo from Wainwright to Wood Buffalo Park 
during the summer of 1924. It was decided that the 
Parks Branch would assume responsibility for segregat- 
ing, corralling and squeeze chuting and placing the buf- 
falo in cars at Wainwright. that the Northwest Territories 
Branch would be responsible for the care and transfer of 
the buffalo from Wainwright to Waterways, thence by 
barge to Wood Buffalo Park. This arrangement was con- 
curred in by the Minister under date of 24th April. 1924. 
Instructions respecting the shipment of the buffalo 
were received by the Superintendent at Wainwright too 
late to permit of the young buffalo being segregated in 
time for shipment in 1924 and it was decided not to 
attempt to transfer any buffalo until the following year. 
The Superintendent of Wainwright Park was given 
instruction to make plans for the transfer of at least two 
thousand young buffalo in 1925. [Note that it is now 
"young" buffalo, not yearlings. 1 

In [al memorandum dated 6th October, 1924. the 
Commissioner of the Canadian National Parks refers to a 
conference which was held in the Deputy Minister's 
office on the 3rd idem when the following decisions 
with respect to the transfer of the buffalo from 
Wainwright to Wood Buffalo Park were made: 

1. In 1925 al least two thousand buffalo would be 
shipped. 

2. The shipment would include only one-and-two- 
year old animals from the 1923 and 1924 increase. 

3. That the tubercular test would be dispensed with. 
[My emphasis] 



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Vol. 116 



4. That the responsibility of the Canadian National 
Parks Branch would cease when the animals were load- 
ed on the cars at Wainwright. 

If we take the Waddy report at face value, that is 
two tubercular animals out of every 15, then around 
900 of the 6673 animals delivered would have been 
carrying tuberculosis. 

The actual ages and numbers transferred over the 
next four years are shown in Table 3. 

Another column in the file listed deaths en route, 
which totalled 47 animals accidentally crushed or 
injured. None of the reported deaths "were attributed 
to tuberculosis." The next sentence, however, reads: 
"The carcasses were not examined to ascertain 
whether the animals were infected, since there was 
no government official available qualified to make 
such an examination." 

My home in Fort Smith in 1947 was no more than 
100 m from the home of an elderly gentleman who 
had been the Chief Warden of Wood Buffalo 
National Park when the transfers were made. He told 
me that many animals died as they walked down the 
gangplank at the landing, or as they made their way 
along a trail that ran for about 2 miles from the river 
through some woods to the nearest patch of prairie. 
Other old timers at Fort Smith, who had taken part in 
the unloading, confirmed the Chief Warden's story. 
When I mentioned this conversation in a letter to 
Ottawa in 1 947 I got a reply that said in no uncertain 
terms that 6673 animals were delivered, apart from 
47 that died en route. Apparently, the translocation 
was still a touchy subject 20 years after it happened. 
The next paragraph in the Richards memo lists the 
names of several people who were opposed to the ven- 
ture. Another document lists arguments pro and con. 

2. Controversy about the translocation 

Two things that caused the most concern among 
opponents of the transfer were first, that infectious 
disease might be introduced to the Wood Bison, and 
second, the Wood Bison would disappear as a sub- 
species through hybridization with the introduced 
plains bison. 

Those who defended the transfer based their argu- 
ments mainly on three untested hypotheses. First 
came Camsell's claim that the northern and the 
southern herds of Wood Bison were separated by a 
band of marsh and muskeg that could not be crossed. 



Thus, regardless of what happened to the southern 
herd, the northern herd would not hybridize and it 
would not be exposed to disease. Second came 
Pike's belief, supported by Graham and others, that 
any differences between plains and Wood Bison 
were superficial and due to environmental factors. 
Some supporters of this view went so far as to pre- 
dict that the introduced animals would gradually 
develop the characteristics of Wood Bison. Third, 
the introduced animals were too young to carry any 
infectious disease including tuberculosis. Both J. B. 
Harkin and Dr. Torrance, however, expressed con- 
cern after the rejection of Dr. Torrance's recommen- 
dation, that only yearlings that passed a tuberculin 
test were to be shipped. 

Among the defenders was Maxwell Graham, even 
though he was no longer associated with the 
Dominion Parks Branch. In a one page Letter to the 
Editor of The Canadian Field-Naturalist (Graham 
1924) he upheld Camsell's belief in the barrier 
between northern and southern herds, and the belief 
that the difference between plains and Wood Buffalo 
was just a matter of different environments. I believe 
that he genuinely believed that the transfer "will be 
the means of saving for posterity the calf crop at the 
Wainwright Park for 1922-23 and succeeding 
years." 

Mr. Edmund Seymour, President of the American 
Bison Association, and Dr. W. T. Hornaday, an 
American mammalogist, met with the Deputy Min- 
ister. The stance taken by the Bison Association 
would have carried considerable weight among the 
general public. However, the Deputy Minister per- 
suaded Seymour and Hornaday to give their approval 
to the actions taken by the government. The warning 
they received in Harkin' s letter, if it really was a 
warning in disguise, was ignored. 

On the other hand, a considerable list of people, 
many of them well versed in mammalogy, expressed 
their objections directly to the government or 
through published articles. 

Dr. D. Rutherford, Department of Agriculture, 
University of Saskatchewan, told the Minister in a 
letter that the transfer was a mistake. 

W. E. Saunders (1925), a naturalist living in 
London Ontario, wrote "it would be better to lose the 
entire Wainwright herd, rather than risk the last rem- 
nant of the Wood Buffalo." 



Table 3. Numbers and Ages of Bison Transported from Wainwright to Wood Buffalo Park 
in the Years 1925 to 1928. 



Season 


Yearlings 


Two Years 


Three Years 


Total 


1925 


1127 


507 


_ 


1634 


1926 


1435 


493 


83 


2011 


1927 


1255 


436 


249 


1940 


1928 


1009 


79 


- 


1088 


TOTAL 


4826 


1515 


332 


6673 



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Professor A. B. Howell, Chairman of the Ameri- 
can Society of Mammalogists, cited a resolution 
passed at their Annual Meeting condemning the 
transfer because "distinctive characteristics of the 
Wood Buffalo would be lost in a few generations," 
and "TB and other diseases would be likely to be 
transmitted with harmful effects to the northern 
herd." 

Dr. Francis Harper (1925: 45), an American mam- 
malogist who had travelled from Lake Athabasca to 
Great Slave Lake with Charles Camsell, and who 
had visited the Park in 1920, made a number of 
points in a letter to the Editor of The Canadian 
Field-Naturalist. First, "The wood buffalo (Bison 
bison athabascae) is too important an animal to be 
subjected to experimentation that may result in 
decided harm to the entire subspecies." Second, 
"The proposal outlined by Graham raises the old 
question of man's interference with nature, which, in 
too many cases is alike unnecessary and unjustifi- 
able." Third, "Establishment of the Park was one of 
the most important and far-sighted conservation 
measures ever adopted by the Dominion Govern- 
ment." The implication is why spoil it? Fourth, with 
respect to the separation of the northern and southern 
herds he wrote "This gap can hardly yet be accepted 
as a proven fact, or as a necessarily permanent con- 
dition. There is evidently no physical barrier that 
would prevent the two herds from mixing." Fifth. 
"The possible transmission of disease through the 
introduced plains buffaloes is another factor to be 
considered." Sixth, if the Wainwrights cannot be set- 
tled in Alberta, "would it not be wiser to send them 
to the slaughter-house at once?" 

Two members of the National Parks Branch faced 
censure over publication of Harper's critical letter to 
the editor. One was Mr. Hoyes Lloyd, Supervisor of 
WildHfe Protection, who was also President of the 
Ottawa Field Naturalist Club at the time. The club's 
journal was, and still is. The Canadian Field Natur- 
alist. The Editor of the journal was Dr. Harrison F. 
Lewis, who later became the first Director of the 
Canadian Wildlife Service. For publishing Harper's 
letter, both men were required to vacate their posi- 
tion in the Club, or face dismissal from the civil ser- 
vice. According to Lothian (1981: 34). Lloyd had 
already written a memorandum to Commissioner 
Harkin in which he called the decision to ship ani- 
mals from a herd known to be infected to a herd that 
had never had contact with the disease, "very bad 
epidemiology," and "the biologically correct way of 



dealing with the excess buffalo is to slaughter the 
excess, thus realizing on the surplus stock." 

Professor William Rowan of the University of 
Alberta visited the Park in 1925 and collected two 
specimens of Wood Buffalo under permit. He pub- 
lished an account of his trip in "Country Life." A 
typescript of part two of his article is included in the 
salvaged files. In his final paragraph he wrote "I 
need not discuss the pros and cons of the proposal 
here, suffice it to say that it called forth, naturally 
enough, the most vigorous protests from scientific 
quarters from all over Canada, from the States and 
even from England." Rowan was black-hsted by the 
Department of the Interior as a result. 

Other condemnations appeared in the two most 
important scientific journals in the world. Dr. 
Ritchie, in England, published his objections in 
Nature. Perhaps the most serious condemnation 
came from Dr. Thomas Barbour. Director of the 
Museum of Comparatives Zoology at Harvard 
University, Boston. In a book review published in 
Science for 25th November 1932, he referred to the 
transfer of bison as follows: 

This, one of the most tragic examples of bureaucratic 
stupidity in all history, was done against the protests of 
both Canadian and American naturalists who would 
rather have seen the surplus bison killed. They were 
known to be infected with bovine tuberculosis and they 
are certain to interbreed as well as infect the wood bison, 
which is a far finer animal and one of great zoological 
interest because in some respects it seems more like the 
European wisent than the common American Bison. 

3. The Aftermath 

Waimvright 

Transplanting a reasonably large number of Bison 
each year did not solve the overpopulation problem 
in Buffalo National Park. As early as the winter of 
1926-27 more or less regular slaughters were 
resumed. There is an accounting of the number of 
Bison killed, and a meticulous record of the money 
received for the sale of Bison products, but there is 
no mention of the prevalence of any disease or para- 
site in the records I have. Table 4 lists the years, and 
the numbers of Bison killed. 

Dr. Seymour Hadwen attended many, but not all 
of the slaughters. The data for Figure 2 come from 
Hadwen (1942). Numbers killed are shown on the 
bars. They are Hadwen' s numbers and some of 
them differ slightly from the government numbers 
in Table 4. 

Hadwen suggested that the prevalence may have 
declined over time. An alternative explanation is that 



Table 4. 


Number of Bison Slaughtered at Wainwright After Transfers Began. 












Year 


1926- 1927- 1929- 1930- 1931- 1932- 1933- 
1927 1928 1930 1931 1932 1933 1934 


1934- 
1935 


1936- 
1937 


1937- 
1938 


1939 


Total 


Number 


2(){)1 lOOO .^2.S 47 1.^34 1220 2000 


1000 


1522 


3246 


29 IS 


17013 



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80.00 T25a 
70.00 



LU 

> 60.00 

g 50.00 

I- 



40.00 



1012 



520 



1522 



15341M2 



^ 30.00 

UJ 20.00 

10.00 

0.00 



POSITIVE FOR T.B. 

Figure 2. Prevalence of tuberculosis in bison at Wainwright (After Hadwen (1942). Numbers over bars are sample sizes. 



2020 




23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 

YEARS AFTER 1900 



the early slaughters concentrated on the oldest ani- 
mals, which were most likely to be positive for 
tuberculosis (Table 2). We know that was true for 
the sample in 1923. It was probably true for 1928 
and 1930, because all three of those years had a 
pre valency rate greater than 70%. If so, as the pro- 
portion of old animals in the slaughters declined, so 
would the prevalence of tuberculosis. 

Wood Buffalo National Park 

Population: The Wainwright bison were unloaded 
either at the Hay Camp, located on the west bank of 
the Slave River about 50 km upstream from Fort 
Fitzgerald, or at Buffalo Landing, which was about 
midway between Hay Camp and Fort Fitzgerald. 
Some accounts mention La Butte as a destination, 
but it is on the east bank of the river, which is not in 
WBNP. Park employees at the time of the introduc- 
tion of the Plains Bison noted that the residents and 
the newcomers mingled almost at once. That does 
not mean, however, that there was a great deal of 
breeding, either with each other, or with the resident 
Wood Bison, in the year of their arrival. Two thirds 
of the total were yearlings, and less than 5% of 
female bison breed as yearlings (Fuller 1962: 21). 

The first attempt to make a census of the Bison in 
WBNP was made in 1931 by the Royal Canadian 
Air Force. They set out to cover the entire winter 
range and take photographs of all herds of Bison. 
Something went wrong because they photographed 
fewer than 2 400 Bison. 

J. Dewey Soper was sent to WBNP in 1932, 
where he stayed until 1934 studying the fauna, espe- 
cially the Bison. He was limited to surface travel — 



canoe and pack horses in summer; dog sled and 
snowshoes in winter. Soper did his best to estimate 
what the numbers might be. His best guess was 
12 000, which is within the realm of reality. 

In 1947, Mr. E. G. Oldham, Superintendent of 
Forests and Wildlife, attempted an aerial survey. 
Instead of attempting full coverage he flew strips so 
placed that he covered one third of the area of the 
park. He counted 2494 bison so his estimate was 7482 
in the park as a whole. Details of his census were in a 
file at Fort Smith when I arrived there in 1947. 

In 1949, I made an aerial strip count based on 
25% coverage (Fuller 1950). My estimate was not 
less than 10 000 or more than 12 500. In a repeat 
aerial census in 1951 (unpublished) I got essentially 
the same result, which suggests that the population 
was not growing, but staying more or less constant. 
All of that came to an end after 1970. 

In 1969 the flow of water in the Peace River was 
cut off near its source when the gates were closed on 
a new dam and the water was held back to fill a reser- 
voir. The flow was "restored" after the reservoir was 
filled. That is to say, about the same amount of water 
comes down the river in a year, but the seasonal dis- 
tribution of the flow is altered. Spring floods are 
essentially a thing of the past. Without spring floods 
every few years the Peace Delta has undergone radi- 
cal change. Perched lakes go dry; what used to be 
navigable small streams running into Lake Clair are 
choked with vegetation; sedges are being replaced by 
grass, and grass by shrubs and trees. All of this has 
had an enormous effect on the number of Bison in the 
part of WBNP south of Peace River (Figure 3). 



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155 



LU 
CQ 



CO 

(O 

z 

LU 
O 




5.500 



12 3 4 5 6 7 8 9 10111213141516171819 20 2122 23 24 25 26 27 28 



YEARS AFTER 1970 



LN (NUMBER) 



REGRESSION 



Figure 3. Natural logarithms of counts of bison south of Peace River beginning in 1971. The straight line is the 
Calculated Regression Line. 



The equation for the regression line for the log^ 
transformed numbers is: LN(Nt) = 9.141 - 0.086t, 
with an r^ of 0.887. The rate of decline over 28 years 
has been 8.6% per year, and the probability that the 
relationship could have occurred by chance is less 
than 0.001. 

Figure 4 shows what has happened to the popula- 
tion as a whole. The striking similarity between 
these two Figures shows that changes in the delta 
have affected not only the southern part of the park 
but the park as a whole. 

The equation for this log^ transformed regression 
is LN(Nt) = 9.137-0.051t, with an r^ of 0.917. The 
overall rate of decline has been 5.1% per year. The 
probability that the relationship was due to chance is 
less than 0.001. 

North of Peace River the population fluctuated up 
and down without any visible pattern. The regression 
line, which actually has a positive slope of 0.5%, is 
meaningless, with an r- < 0.001. 

Tuberculosis: Col. Richards ended his summary of 
correspondence concerning the transfer with two 
short, but interesting, paragraphs. 

The information available lends support to the view 
that as only young animals were being transferred to the 
Wood Buffalo Park, improvement in their condition 
might be looked for. It is doubtful whether any of our 
park officials are in a position to definitely determine to 
what extent the animals in Wood Buffalo Park are 
affected with tuberculosis. In these circumstances it 
would seem that facts are not available to justify the 
conclusion that tuberculosis is prevalent or endangering 
the buffalo in Wood Buffalo Park. 



It is the practice to authorize the killing of a number 
of aged buffalo in Wood Buffalo Park each year, the 
meat of which is distributed to charitable institutions and 
to natives and half-breeds in the northern districts. In 
view of Mr. Soper's report it would seem that the ser- 
vices of a veterinarian should be available at the time of 
the buffalo slaughter to ensure that the meat of healthy 
animals only is being disposed of for the purposes men- 
tioned. 

When I arrived in Fort Smith in 1947 the term was 
'buffalo hunt' not slaughter. The hunt began in the 
fall after it got cold enough to preserve the meat. Old 
males were the targets even though it had been 
known since the 1923 slaughter at Wainwright that 
the prevalence of tuberculosis was highest in aged 
animals, especially males. Park staff did all the 
shooting. Skinning, butchering, and transporting 
were the responsibility of the recipients. The Fort 
Smith hunt took place on the Salt Plains, which 
could be reached by 4-wheel drive vehicles. The Fort 
Chipewyan hunt took place on part of the Peace- 
Athabasca Delta where the only means of transporta- 
tion was the dog sled, and the whole crew slept in a 
two-room cabin. 

In 1950 a professional butcher and a veterinary 
meat inspector were brought from Edmonton for the 
hunt at Fort Smith. I worked with the veterinarian, 
who showed me where to look for signs of tubercu- 
losis, and what the signs are. I carried on as "inspec- 
tor" for the animals killed at Chipewyan. Thus ended 
about 25 years of feeding uninspected carcasses to 
school children and people on welfare. 



156 



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UJ 
CD 

ID 
Z 

CO 

3 
CO 

z 

LU 
O 



9.400 



9.200 




12345678 9 10111213141516171819 20 2122 23 24 25 26 27 28 

NUMBER OF YEARS (-1-1970) 

■ LN(NUMBER) -^ REGRESSION 

Figure 4. Natural logarithms of counts of bison in Wood Buffalo National Park as a whole beginning in 1971. The 
straight line is the Calculated Regression Line. 



For two or three years a mobile "abattoir" was 
used. It consisted of three buildings on sleds pulled 
by a tractor. Finally, permanent abattoirs were con- 
structed at Hay Camp and on the Peace Delta close 
to Sweetgrass River. A Veterinarian was present at 
all slaughters once the abattoirs were in use. 

As an example of the findings. Figure 5 shows the 
prevalence of tuberculosis in 527 males and 981 
females examined at Hay Camp in the years 
1952-56 (Fuller 1962: 29). Age Class means calf, 
Classes 1 to 4 are ages 1 year to 4 years, Class 5 is 
Young Adult, Class 6 is Prime Adult, and Class 7 is 
Old Age. Characteristics used to define each age 
class are set out in Fuller (1959). 

It is clear from Figure 5 that the prevalence of 
tuberculosis increased with age as it did at 
Wainwright. Almost three-quarters of adult and old 
males were positive. Overall, 38 per cent of males 
and 40 per cent of females showed evidence of 
tuberculosis. There can be no denying that tuberculo- 
sis had been introduced to Bison in WBNP. 

Brucellosis (Contagious Abortion): The origin of 
Brucellosis in the park is a mystery, but it is most 
likely to be the Wainwright animals. Brucellosis was 
first confirmed in the park in 1956, which was the 
year that I left Ft. Smith. Some symptoms character- 
istic of Brucellosis had been seen during slaughters, 
but laboratory confirmation was lacking because it 
had not been possible to keep samples from freezing 



in the camps and on their way to the lab. The preva- 
lence of Brucellosis at the Sweetgrass abattoir during 
its period of use (1957-1958 to 1973-1974) was 
39.5%. A total of 1681 individuals were sampled of 
which 664 were positive. 

The incidence of tuberculosis at Hay Camp during 
the same period was 31%. Given these two results, 
the probability that an animal chosen randomly did 
not have at least one of the two diseases was 0.417 
or about 4 chances out of 10. 

Recent studies of Bison diseases were reported by 
Dr. Stacey Tessaro (1989) and Tessaro et al. (1990). 
Tessaro (1989:5) found "...brucellosis in 18 (25%), 
and tuberculosis in 15 (21%), of 72 Bison that had 
died as result of hunting activity, wolf predation, nat- 
ural accidents, or disease. The combined prevalence 
of the two diseases in the sample was 42%. Of the 56 
Bison that were killed, butchered and utilized by 
native hunters, 24 (43%) had one or both of these 
two zoonotic diseases. Three or four Bison seen 
killed by wolves were severely debilitated by gener- 
alized tuberculosis, and I suspect that a significant 
portion of the adult Bison mortality attributed to 
Wolves is predisposed by disease. Another cow was 
found to have died of pulmonary tuberculosis. 
Brucellar arthritis in three Bison resulted in crip- 
pling, emaciation and death of these animals. Eight 
other severely lame and emaciated bison were seen 
in the park, but these animals could not be collected 
for scientific evaluation." Tessaro also expressed 



2002 



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157 





80.00 




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> 


60.00 


H 




(0 


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o 


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ct 




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3 4 

AGE CLASS 

■ MALES ■ FEMALES 

Figure 5. Prevalence of Tuberculosis by sex and age group in 1508 bison examined at Hay Camp, WBNP, in the years 
1952-1956. (After Fuller, 1962). Numbers above bars are sample sizes. 



concern that the diseases could be carried to the 
Mackenzie Bison Sanctuary by one or more roaming 
park bison. 

Mackenzie Bison Sanctuary 

In 1958, Dr. N. S. Novakowski located a group of 
about 200 bison with many of the characteristics of 
Wood Bison near the northern boundary of the park. 
In the next few years many of them were trapped and 
moved to corrals set up on the Salt Plains near Fort 
Smith. There they were tested repeatedly for tubercu- 
losis and brucellosis. Reactors were eliminated after 
each test. Drs. Banfield and Novakowski (1960) 
examined skeletal material as well as characteristics 
of the living animals, and declared that the bones and 
the live animals belonged to the Wood Bison sub- 
species (Bison bison athabascae). Their opinion has 
been both questioned and supported several times 
since they published the results of their study. This is 
not the place in which to debate taxonomy, but there 
is no question that those animals and their descen- 
dants are the closest we will ever see to the original 
Wood Bison. In 1963, the captives at Ft. Smith were 
deemed disease-free. Eighteen of them were moved 
to Ft. Providence where they were released in an area 
north of the Mackenzie River. The area set aside for 
them is known as the Mackenzie Bison Sanctuary 
(MBS). The remainder were sent to Elk Island 
National Park. For more information about their fate 
seeC. C.Gates etal.(2001). 



The little group translocated to the MBS now 
numbers in the region of 2600 head (N.S. 
Novakowski, personal communication). No road 
runs through the MBS, but the highway between 
Yellowknife and Fort Providence runs along its 
western boundary and travellers frequently see Bison 
on or beside the road. 

Although the Mackenzie River separates the MBS 
herd from WBNP, the river is not an impenetrable 
barrier. It can easily be crossed on the ice once it has 
frozen over. The probability of a diseased animal 
from Wood Buffalo National Park wandering as far 
as the MBS is low but non-zero. The herd, however, 
is too valuable to be exposed to the slightest danger 
if it can be prevented. 

Conclusions 

My first objective in this paper has been to show 
that, clearly, it can be argued that the Canadian gov- 
ernment made a serious mistake when it gave in to 
public pressure some 75 years ago. Buffalo National 
Park was bursting at the seams with Bison. Not only 
was the range threatened by overpopulation, but a 
significantly high prevalence of tuberculosis had 
been demonstrated in 1923 by Hadwen and by the 
Waddy Report in 1924. The government of the time 
had at least three choices. One was to continue to 
slaughter large numbers every year in order to keep 
the population within the carrying capacity of the 



158 



The Canadian Field-Naturalist 



Vol. 116 



range. The second was to move surplus animals to 
the new WBNP and take a chance that the diseases 
would be left behind, and that at least some of the 
Wood Bison would avoid contact with the intro- 
duced Plains Bison. The third choice, which was 
proposed by Hadwen, and backed up by Dr. Clarke, 
was to slaughter the entire population, allow the 
range to recover for a few years, and restock Buffalo 
National Park with healthy animals that were already 
available in Elk Island National Park. The third 
option was the only one that guaranteed that no dis- 
eases would be introduced to the Wood Bison, nor 
would any hybridization between Plains and Wood 
Bison take place. 

Hewitt's suggestion to sell surplus Bison to farm- 
ers faced several difficulties. Among the most seri- 
ous was that the Park lacked the facilities for holding 
and testing hundreds of animals every year in spite 
of Graham's repeated pleading for the necessary 
equipment. It would have been highly unethical, and 
probably illegal, to knowingly sell diseased animals 
that might end up on the farmer's dinner table. 

The government put its faith in the second option, 
which rested on the three untested and unproven 
hypotheses set out in this paper. The test of those 
hypotheses was the transfer itself and, as we have 
seen, all three hypotheses were untrue. Unfor- 
tunately, the experiment was non-reversible. 

My second objective in preparing this paper was 
to discharge the obligation I assumed regarding 
Maxwell Graham. I believe that the excerpts from 
the old files show that Graham was devoted to the 
Bison at Wainwright. He showed continuous con- 
cern for their health and was devastated when he 
learned that tuberculosis was present in the herd. He 
was cleared of all blame by two of his superiors in 
the civil service, one of whom was the Deputy 
Minister. He played no role in the decision to trans- 
fer young bison to the newly formed Wood Buffalo 
National park, and he had left the National Parks 
Branch some two years before the transfer began. 
His one transgression was the one-page letter he had 
published in The Canadian Field Naturalist that sup- 
ported the three failed hypotheses. We will probably 
never know whether he wrote that letter on his own 
initiative or whether he was "asked" by one of his 
superiors to write it. There is no doubt that the annu- 
al calf crop in the mid 1920s at Wainwright faced an 
uncertain future, and I can understand why Graham 
sincerely believed that the transfer gave the young 
animals their best chance for survival. 

In August of 1990 the Report of the 
Environmental Panel on Northern Diseased Bison 
was made public. The recommendation of the Panel, 
of which I was a member, was that all the Bison in 
Wood Buffalo National Park should be slaughtered. 
During and after the slaughter. Bison from Elk 
Island National Park should be bred in small enclo- 



sures in the north so that restocking with disease-free 
animals could take place as early as possible. The 
recommendation met with opposition on many fronts 
and, shades of 1923, the government backed down 
once again. The government decided to "study" the 
situation. Ten years on, no action has been taken. 

So here we are in the new Millennium, and we are 
faced once again with a situation similar in many 
respects to that faced by the managers of Buffalo 
National Park in 1923-24. There is a herd of bison 
with a high prevalence of two infectious diseases 
and a declining population. There is another herd 
that is disease-free and flourishing. There is a low, 
but non-zero probability of infection of the clean 
herd by diseased animals wandering from their prop- 
er home. The Panel recommended total slaughter 
and replacement, but it did not recommend slaughter 
just for the sake of killing. 

We must keep in mind what happened at Wain- 
wright when the government refused to do a com- 
plete slaughter after it was shown that up to 75% of 
old animals were infected with tuberculosis. In 1923, 
the Bison population was under 7000. The total loss 
of life would have been less than 7000 if the whole 
herd had been slaughtered as Dr. Hadwen recom- 
mended. Ultimately, the protesters and a weak gov- 
ernment were responsible for the deaths of 17 000 
Wainwright Bison (Table 4), for introduction of dis- 
eases to the pristine population of Wood Bison, and 
for contaminating the majestic Wood Bison by 
hybridization with Plains Bison. 

The situation in WBNP, unlike that in Wainwright, 
has actually improved with time. In 1990, the Bison 
population was more than 3300. In 1998, it was just 
over 2300 and declining. Some of the annual slaugh- 
ters at Wainwright were as large as, or larger than, 
what would be needed for a final clean-up in WBNP. 

There is no logical reason for further delay. 

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Cameron, A. E. 1924. Some further notes on buffalo. The 
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tional range: The threat of tuberculosis and brucellosis. 
Wildlife Veterinary Report. 2: 5-6 

Tessaro, S. V., L. B. Forbes, and C. Turcotte. 1990. A 
survey of brucellosis and tuberculosis in bison in and 
around wood buffalo National Park, Canada. Canadian 
Veterinary Journal. 31: 174-180. 

Accepted 12 April 2001 
Received 1 1 March 2002 



Book Reviews 



Zoology 

Birds, Mammals & Reptiles of the Galapagos Islands 



By Andy Swash and Rob Still. 2000. Pica Press and 

WildGuides, Yale University Press, New Haven and 
London. 168 pp., illus. U.S. $24.95. 

I was delighted to find a sturdy, handy-sized guide 
to all the animals of the Galapagos Islands. For the 
traveler, size as well as quality is important. This 
book certainly is the right size and packs all the 
essential information into one volume. It covers all 
of the resident mammals, reptiles, and birds. It also 
covers the migrant and vagrant birds as well. There 
are introductory sections on geography, climate and 
habitat. 

So the size and content are good, but what of the 
quality of the information? This is somewhat of a 
mixed bag. The introductory sections are logical, 
concise, and clear. They contain as much as I need to 
have in a book I plan to carry in my pocket while 
trudging across a tropical island. The bird section 
begins with an introduction to the major bird groups. 
The authors have used a modified taxonomic group- 
ing that works well. This leads into the main section 
on the bird species. 

This book relies on computer-manipulated digital 
images as the main form of illustration. As a result the 
plates look more like a "standard" field guide of the 
Peterson type. This is an improvement over the old 
sequence of small pictures. However, my arguments 
against photographs in field identifications still hold. 
For example, the Black Petrel and the Hoary Bat have 
a strong rosy tinge, giving a distorted impression of 
their true colours. The juvenile Yellow Warbler has a 
powder blue head. Several other birds had odd 
colours, including lilac-toned Lava Gull and Sora. ^he 
strangest transformation is on the female Belted 
Kingfisher. The breast bar is clearly visible, but the 
belly bar is so obscure a novice might miss it. Yet 
more strange is the breast bar is distinctly chestnut, 
instead of gray. Is this computer manipulation gone 
awry? Other birds on the same page as the odd ones 
were normally coloured. I also wonder why the 
authors did not change the backgrounds to give more 
contrast with the birds. They have brownish herons on 
a brownish background; grayish mockingbirds on a 
grayish background, and so on. If they wanted to 
manipulate why not follow their own fine example, 
the flycatcher page. This shows the birds clearly and 
in good proportion. 



The text explains that the light morph of the 
Wedge-tailed Shearwater is more likely yet the plate 
depicts the dark morph. I would also prefer that the 
authors had emphasized the flight of oceanodrama 
petrels, a much more useful characteristic in the 
field. 

One plate is devoted to the heads and beaks of 
Darwin Finches. The heads are depicted against a 
graph paper background and the outline of each bill 
is picked out in an unobtrusive gold line. Coupled 
with the description this is a useful tool for the inex- 
perienced visitor. 

The whale section is, for the most part, well done, 
with the species being depicted as you see them from 
a ship. While this does not show much of each 
species, it does present a realistic view of a typical 
field sighting. It also demonstrates the difficulties of 
field identification. I was not happy with the photo- 
graph of Cuvier's Beaked Whale. It was dark and 
showed little of this species' characteristics. The 
authors did mention the possible spotting on a Sei 
Whale. The Blainville's Beaked Whale is coloured 
red. I have not seen a Blainville's Beaked Whale, but 
the other beaked whales I have seen were at best 
described as brownish. I would be surprised to find a 
maple leaf red specimen. 

I would like to see the authors add information on 
seasonal distribution. I realize most of the birds are 
year round residents, but there are others I question. 
Are the "resident" Waved A present all year or they 
disperse at the end of their breeding season? This 
information would also be valuable with scarce 
migrants. When can we expect to see skuas or 
jaegers? 

Despite my comments, this is a useful field book. 
It is unlikely that anyone would confuse any of the 
species, despite the odd errors the species that could 
be confused are well covered. The book covers all 
the Galapagos animals in one handy portable vol- 
ume. So I will try it in the field in late October this 
year. In the meantime I will be using other higher 
quality, but far less portable, texts for research. 

Roy John 



2193 Emard Crescent, Beacon Hill North, Gloucester, 
Ontario KIJ 6K5 Canada 



160 



2002 



Book Reviews 



161 



Wisconsin Fishes 2000: Status and Distribution 

By John Lyons, Philip A. Cochran, and Don Fago. 
University of Wisconsin Sea Grant Publication 
No.WISCU-B-00-001. University of Wisconsin Sea 
Grant Institute, 1975 Willow Drive, Second Floor, 
Madison, Wisconsin 53706-117. e-mail: linda@sea- 
grant.wisc.edu. 87 pp. U.S. $10. 

One wonders if the Electronic Age will make peer 
book reviews less important, or more important. An 
eight-page description of this book is available at 
http://www.seagrant.wisc.edU/greatlakesfish/Lyons.h 
tmf. This is my first experience evaluating a publica- 
tion with most of the routine items, or samples, read- 
ily at hand. 

The header on each of the eight pages of the 
Internet description uses the statement "Wisconsin 
Fishes 2000/Fish of the Great Lakes by Wisconsin 
Sea Grant." This Great Lakes emphasis is not men- 
tioned in the book. Great emphasis is placed, by the 
authors, on the literature since Becker's (1983) 
Fishes of Wisconsin, and the fact that this publica- 
tion updates the information on "occurrence, taxo- 
nomic status, and abundance of fishes in Wisconsin". 
Fishes in the waters of the Wisconsin portions, and 
at times the adjacent Michigan portions, of lakes 
Michigan and Superior are, however, included. 

Individuals interested in the status and distribution 
of fishes in Wisconsin, and comparisons with those 
in the geographic areas around Wisconsin in the 
USA and Canada, owe a debt of thanks for the 
extensive and constantly updated information since 
Becker's 1983 book, in the 22 related publications 
by these authors and coauthors, and those of several 
other authors. Much of the information in this book 
was derived from the "Master Fish File", a database 
including 22 000 Wisconsin fish collections from 
1900 to 1999. The database can be accessed through 
the web site of the Wisconsin Department of Natural 
Resources (www.dnr.state.wi.us). 

This book includes sections entitled Summary, 
Introduction, Materials and Methods, Overview of 
Changes in the Wisconsin Fish Fauna, Species 
Accounts, References, Index to Common Name, 
Index to Species by Scientific Name, and eight 
colour plates. 

The overview of changes includes a table with 
sections entitled Native Species (147), Established 
Non-Native Species (14), and Transient Non-Native 
Species (19). In total those sections include 180 
species. One of the problems of lists is the neces- 
sary, sometimes arbitrary, decisions required when 
authors decide where to put various species. Readers 
may not have made the same decisions. 

Five species not included in earlier publications are 
included here. Those are Kokanee, Oncorhynchus 
nerka, Threespine Stickleback, Gasterosteus aculea- 
tus. White Perch, Morone americana, and Round 
Goby Neogohius melanostomus. 



There are species accounts for 182 forms, ranging 
in length from a few lines to several pages. The 
emphasis, in length, is on the more recently discov- 
ered species, native and introduced. Many of those 
were not given full treatment in Becker's 1983 book. 
Only eight of the species accounts include a "spot" 
map of those locations at which the species has been 
collected, or observed by one of the authors. The 
longer species accounts usually include Common 
Name; Scientific Name; Description, including 
means of separation from look-alike species; 
Systematic Notes; Distribution, Status, and Habitat; 
Biology; and Importance and Management. The dis- 
tribution statements, and absence of a map, in the 
shorter Species Accounts leaves the reader unable to 
determine if the species occurs in lakes Michigan 
and/or Superior. Some species accounts deal with 
extirpated species, and species rediscovered as pre- 
sent. 

The book includes eight colour plates tipped into 
the front before the title page, somewhat like a 
Frontispiece. The plates are of seven recently dis- 
covered forms. There is one unfortunate problem. 
The caption of one of the two plates for the Round 
Goby, Neogobius melanostomus, incorrectly identi- 
fies the "fused pectoral fins", whereas the photo 
clearly indicates that it is the pelvic fins which are 
fused forming the diagnostic "sucking disc" of gob- 
ies. 

There are a number of interesting points to be 
derived from the lists and accounts. The abundance 
of natural reproduction of Lake Trout, Salvelinus 
namaycush in the Wisconsin portion of Lake 
Superior has led to stocking being discontinued 
there. The same is apparently true for Chinook and 
Coho salmon, Oncorhynchus tshawytscha and O. 
kisutch. In contrast, for Pink Salmon, O. gorbuscho, 
there has been no natural spawning in Lake Superior 
streams since the late 1970s and the species is now 
rarely seen in Wisconsin waters of lakes Superior 
and Michigan. There have apparently been no 
records of the European Flatfish, Plotichthys flesus 
from Wisconsin waters of lakes Michigan or 
Superior. This is in contrast to the fact that Ontario 
records for Lake Superior are almost as numerous as 
those for Lake Erie. The Warinouth, Lepomis gulo- 
sus has now been reported from (the southern part of 
?) the Lake Superior Basin. This species is easily 
confused with the Rockbass, Ambloplites mpestris, 
and even though Lake Superior may be more of a 
barrier (cold) to immigration, with the experience on 
Lake Erie its arrival in Lake Superior should be 
watched for. 

There is, presently, a great emphasis on mainte- 
nance of biodiversity, rehabilitation of native 
species, and the controversy aroinid "Is more bet- 
ter?". This commitment cannot hope to succeed 



162 



The Canadian Field-Naturalist 



Vol. 116 



without constant surveys, species status reports, 
checklists, and publications like this one and the oth- 
ers which have appeared in Wisconsin. Too often, 
surveys and updates, such as this one, are deemed 
less important than "more sophisticated" research. I 
hope the people in Wisconsin will continue to be 
supported philosophically and financially to set an 
example for other Political areas. I found the book 
interesting and useful. I'm certain that fish biologists 
in Wisconsin are finding it useful, as will the next 
person charged with updating a checklist of the fish- 
es of the Great Lakes Basin. 



References 

Becker, G. C. 1983. Fishes of Wisconsin. University of Wisconsin 
Press, Madison, Wisconsin. 

Robins, C. R.,R. M. Bailey, C. E. Bond, Jr., Brooker, E. A. 
Lachner, R. N. Lea, and W. B. Scott. 199L Common and sci- 
entific names of fishes from the United States and Canada. Fifth 
Edition, American Fisheries Society Special Publication 20, 
Bethesda, Maryland. 

E. J. Grossman 

Curator Emeritus, Ichthyology, Centre for Biodiversity and 
Conservation Biology, Royal Ontario Museum, Toronto, 
Ontario M5S 2C6 Canada 



Marine Mammals of the Pacific Northwest: A Concise and Comprehensive Waterproof 
Guide 



By P. A. Folkens. 2001. Harbour Publishing, Madeira 
Park, British Columbia. 8 pp., illus. $9.95. 

The increasing development of the whale-watch- 
ing industry has resulted in the demand and creation 
of books and identification guides for sea mammals, 
seabirds, and other parts of the marine ecosystem. In 
this regard, and looking back on a history of whale 
watching for over 90 years, the coast of British 
Columbia has received most of the publication activ- 
ity in Canada. 

Despite its catchy title, this guide by Pieter Arend 
Folkens is more a leaflet than a guide. It consists of 
three text pages and five pages of drawings and pho- 
tos full of information for the whale watcher in the 
field. Although the leaflet is printed on "waterproof, 
UV resistant synthetic film made from a 100% recy- 
clable, environmentally inert material containing no 
forest products (similar to milk jugs)", the user 
might actually have difficulties using it on an off- 
shore whale watching trip during periods of strong 
wind or high waves; the light leaflet could easily fly 
away and the small print is hard to read when on a 
rolling boat. However, the compressed text gives a 
nice summary and overview on 31 sea mammal 
species in the area; it even mentions Steller's Sea 
Cow which was hunted to extinction by 1768. In 
addition, major whale watching locations in British 
Columbia and Alaska are named; but none are 
specifically reported for Oregon and Washington (as 
the title would imply). 

Most of the eight pages of the field guide are devot- 
ed to drawings and to fine pictures from the author and 
several others. The reader might find the distinction 
between Mysticetes (Baleen Whales), Odontocetes 
(Toothed Whales), and Small Cetaceans a little unclear 
from the arrangements of the drawings. Very helpful 
and informative is the page about "Common visible 
behaviors and terms" allowing to link sea mammal 
sightings to a classified set of behaviour types. Helpful 
also is that images of fiuke displays are presented for 
species that are known to show such behaviour. Even 



the body sizes of newborn sea mammals are given. All 
measurements are made in SI units, and the conversion 
factor for feet is provided. 

Of interest to the general audience might be the 
section "Marine Mammal Watching Guideline," also 
presented on the web [http://www.fakr.noaa.gov/ 
protectedresources/mmvie wingguide.html] (Note 
that the old URL www.nmfs.gov/prot_res.html and 
given in the guide was updated). In addition, 
contact addresses and a web address (revised to 
http://www.fakr.noaa.gov/protectedresources/strand- 
ings.htm) are given for sightings of stranded sea 
mammals (Canada: Department of Fisheries and 
Oceans 800 465-4336; or the Whale Reporting & 
Stranding Line 800 665-5939). 

As in many other field guides, the text suggests 
some field marks and details for species identifica- 
tion and separation that normal whale watchers 
might not be able to apply, or which are not really 
realistic. For instance, Sei Whales are supposed to be 
differentiated from Fin Whales by a fin angle of over 
45 degrees; male Beaked Whales (Genus Meso- 
plodon) ideally can be identified by the location of 
teeth and jaw line (which is, for most of the time, 
covered by the ocean); phocids (true seals) differ 
from otariids (Sea Lions and Fur Seals) by their hair 
and small nails on their foreflippers. Overall, fea- 
tures like these might be very hard to recognize for 
the untrained as well as for the trained observer, par- 
ticularly when observations are made of moving ani- 
mals some distance from a shaky boat, with binocu- 
lars. Rather than focusing on classical small-scale 
features, an outline of the use of proportions and 
shapes could be more useful for distinguishing 
species. Counterproductive for a field guide might be 
the point that Beaked Whales, the species group that 
lacks most knowledge on distribution and where 
whale watchers could indeed contribute greatly to 
science, are described as the "most difficult whales 1 
to identify correct." No further help or details are ■ 
given for the interested whale watcher. 



J 



2002 



Book Reviews 



163 



For my taste, the "Habitat and Symbol Keys" that 
are supposed "to narrow possibilities in a particular 
era" and link sea mammals with "habitat" are not 
really helpful. Many whales migrate across habitats 
and the regular observer has no real way to tell "tem- 
perate" habitats apart from "cool temperate" ones. 
The meaning of the orange W habitat class presented 
for the False Killer Whale will likely remain a mys- 
tery to the reader because its meaning is unexplained 
in the guide. For pinnipeds, their "calls" and the 
mention of rookeries could have been helpful. 



Overall, it appears that this "guide" is an excerpt 
of a better and larger guide book from the same 
author. It is useable in the field, but does not replace 
the real and classical guide books. 

Falk Huettmann 



Centre for Wildlife Ecology, Biology Department, Simon 
Eraser University, 8888 University Drive, Bumaby. British 
Columbia V5A 1S6 Canada. 



Manitoba Birds 

By Andy Bezener and Ken de Smet. 2000. Lone Pine 
Publishing, Edmonton, Alberta. 176 pp., illus. $17.95. 

I often wonder in which market book writers think 
they will sell their product. My first reaction to this 
compact little book, covering only 145 species (out 
of the official list of 406) was that it was too simple 
for most birdwatchers. And it probably is. Then I 
realized it was ideal for scout leaders. It gives a 
short, easy-to-read account of a bird's character, and 
a sentence or two on identification, size, status, habi- 
tat, nesting, feeding, voice, similar species, and the 
best sites in the province (to see the bird). An illus- 
tration and a range map accompany this information. 
This is just about the right level of detail for a scout 
leader teaching his troop about birds. It is also good 
for schoolteachers, new birders, and children. While 
it is intended for use in Manitoba, it would also be a 
useful guide throughout the prairies. 

The introduction has a map of the natural regions 
showing the best 50 birding sites. A dozen sites have 
a short write-up that includes the most prominent 
bird species. I liked this addition to the text. I 
thought beginners and visitors would be able to use 
it as a starting point when organizing trips. Each 
account has a small illustration. While most of these 
are reduced (and reversed) versions from the main 
text, five species are different and not covered in the 
core section of the book. I counted 145 in the main 
text, so there is a total 150 species illustrated. These 
additional species, which this guide explains you can 
find most easily in the places mentioned, would be 
better included in the central body of the book (in 
replacement for some of the rarer or more difficult to 
see birds). 

The authors claim the birds selected are the most 
common and easily found within the province. I 
question some of the choices. They have chosen, for 



example. Burrowing Owl, a rare nester in the south 
west comer of the province and Screech Owl which 
nests along a narrow strip close to the U.S border 
(the range map for this owl seems a trifle optimistic). 
Yet they left out Hawk Owl, a widespread bird 
which nests over the northern three-quarters of the 
province. Birds seen easily in the north (Churchill) 
but not in the south, such as Bonaparte's Gull, 
jaegers, ptarmigan, and plovers, are also not includ- 
ed. Generally, though, I found the choices to be rea- 
sonable for the populated south of the province. I 
would be surprised if House Sparrow is not found at 
northern towns like Lynn Lake and Thompson (the 
range map does not suggest this.) 

The illustrations by Gary Ross, Ted Nordhagen, 
and Ewa Pluciennik are, for the most part, very 
good. They generally show a typical adult in summer 
plumage. For some, but not all, species the differ- 
ences between male and female are shown. Winter 
visitors are depicted in the appropriate non-breeding 
plumage. Many birds are shown in flight too. I was 
very impressed with the shape, attitude, and techni- 
cal detail of each species. I did find a few that I 
thought could be improved (Warbling Vireo is too 
pale and Ohve-sided Flycatcher is not heavily built 
or boldly coloured enough), but overall the quality is 
first rate. 

This is a compact (14 X 21 X 1 cm) little book 
that will easily slip into a pocket or pack. The plasti- 
cized cover and high quality production suggests it 
should last well in the field. A quick colour-coded 
guide to the bird families on the back cover will also 
help out novices to find the appropriate text. 

Roy John 

2193 Emard Crescent. Beacon Hill North. Gloucester. 
Ontario KIJ6K5 Canada 



164 



The Canadian Field-Naturalist 



Vol. 116 



Heron Conservation 

Edited by James A. Kushlan and Heinz Hafner. 2000. 
Academic Press, London and San Diego, xvi + 480 pp. 
illus. 

This publication is a cooperative one, with 19 
major contributors and a multitude of cooperators 
from the Herons Specialist Group, an international 
network of experts in the field of herons and wetland 
protection. Their objective was "to synthesize and 
summarize the state of knowledge of the conserva- 
tion needs of herons throughout the world." 

The book opens with eight chapters on the status 
of herons in major geographic regions — either con- 
tinents or sub-continents. The herons of each area 
are reviewed species by species, with estimates of 
their numbers, or at least their status and conserva- 
tion needs. Of necessity there is much variation in 
the coverage, with considerable detail available on 
European species and lamentably little for many 
Asian and South American ones. Next come seven 
chapters on various aspects of heron conservation, 
covering such topics as habitat conservation, aqua- 
culture, and environmental contaminants, then a 
chapter on research and information needs, and final- 
ly a synthesis of the conservation needs for the most 
vulnerable species. There is an extensive bibliogra- 
phy and a well-organized index. 

Any book devoted to the conservation needs of 
any group of species on a global scale these days 
makes for depressing reading, and this work is no 
exception. Information is lacking on even the most 
basic aspects of life history and distribution of many 
more elusive herons, and the sad chronicle of habitat 
loss, exploitation, and persecution is common to all 
too many species cross the world. 

Still, this is hardly news to anyone interested in 
birds, and I found myself wondering about the audi- 
ence for a work of this kind. Action on any individu- 
al species or group of species will necessarily be 
quite specific to the local conditions and the needs of 
the birds. Much of the information given here, how- 
ever, is very general, perhaps inevitably so given the 
enormous scope of the book. One would think this 
information would already be well known to those 
working in the field, yet at the same time the very 
specific scope of the book limits its appeal to a 
broader audience. 

In an attempt to assess its relevance to those 
species that I am most familiar with, I placed partic- 
ular emphasis on the North American chapter. I was 
left with a distinct sense of disquiet. Its general tone 
of optimism runs contrary to my own experience on 
an admittedly very limited, local scale. It's true, of 
course, that our problems are minor when compared 
to the enormous challenges elsewhere, but at the 
same time the problems are real. I'm not sure that 
cheerfully quoting the Canadian government's poli- 
cy of "no net loss" of wetlands is particularly helpful 



when — as the authors themselves concede — it 
only applies to the 29% of wetlands that are federal- 
ly owned, most of which are north of significant 
heron range. Many of the wetlands I know are pri- 
vately or provincially owned, and dying the death of 
a thousand tiny cuts, while supposedly safeguarded 
by underfunded agencies reporting to unsympathetic 
governments. I question the statement that "It is like- 
ly that, overall, the loss of wetland has been halted in 
Canada . . ." 

I found myself equally at odds when considering 
some individual species, again finding more seeming 
optimism than I can share. On the Least Bittern, for 
example, the authors do stress the lack of good data 
and the fact that in 1986 birders were already 
expressing concerns about the species. But then the 
1965-1979 Breeding Bird Surveys, of all things, are 
cited as indicating "some evidence" of increases. 
The reference is correct as far as it goes, but when 
one considers that in 1997, for example, a grand total 
of 18 Least Bitterns were recorded on only seven of 
1832 BBS routes continent- wide, one wonders how 
useful its' information is in the present context. 

But perhaps the authors are indeed right on a con- 
tinental scale, and then the bothersome part is their 
conclusions could easily seem to justify complacen- 
cy generally. They would doubtless respond that all 
the caveats about lack of data and "alteration of wet- 
land functions" [replacement of marshes with boat 
docks, perhaps?] are there. Indeed they are, but they 
only "temper" the rosy picture they have already 
drawn, poor data or not. 

However, none of the other continental authors 
seem quite so comfortable with the conditions their 
herons face, and the book as a whole seems to 
achieve its objective of pinpointing those areas that 
need action to conserve heron species world-wide, 
and the kinds of action needed. The chapters on 
research and information needs, and the final synthe- 
sis should be particularly useful in this regard. If one 
issue emerges as of paramount importance every- 
where as a critical first step, it is the pressing need 
for more data, and particularly more precise data, to 
allow species' status to be understood. 

This book is an important one for anyone interest- 
ed in heron conservation. While North American 
readers may share my irritation at the authors' 
approach, careful reading will still yield a reasonable 
summary of the issues we face in conserving our 
herons. However, the book's appeal to a broader 
audience is necessarily limited. 

Clive E. Goodwin 



1 Queen Street, Suite 401, Cobourg, Ontario K9A IMS 
Canada 



2002 



Book Reviews 



165 



The Field Guide to the Birds of Australia 

By G. Pizzey and P. Knight. 1999. Revised Edition. Angus 
& Robertson, Sydney, Australia. 576 pp., illus. AUD 
$39.95. 

The success or failure of any field guide depends 
on its ability to convey information necessary for 
correct species identification. The latest edition of 
The Field Guide to the Birds of Australia by veteran 
author Graham Pizzey and illustrator Frank Knight 
accomplishes this objective, and does so in an 
appealing format. 

The introduction to this edition provides a primer 
for those new to birding, or for those wishing to 
"brush up" on their terminology. A generous mix of 
illustrations, text, and examples are used to familiar- 
ize the reader with descriptions of anatomy, field 
marks, and behaviours. Although reading this section 
is not essential to understand the field guide, it will 
assist the novice birder in gaining the most from a 
birding excursion. 

Following the introduction, there are species 
accounts for 778 native and introduced species with 
700 distribution maps. Typically, there are 3-4 
species accounts per double page with text and color 
illustrations opposite each other. Descriptions for 
each species include information on (1) other names, 
(2) similar species, (3) voice, (4) habitat, (5) breed- 
ing season, (6) nests, (7) eggs, and (8) range and sta- 
tus. The range information is particularly useful 
since it refers to place names, easily found on maps 
conveniently included inside both covers. The colour 



illustrations are well done and depict male, female, 
and juvenile plumage; races and moult patterns also 
are provided to facilitate identification. 

The species accounts are followed by an introduc- 
tion to the bird families of Australia. General charac- 
teristics of each family (i.e., behaviour, food, and 
range) are presented in the same order in which they 
appear in the species accounts. The world range of 
each family is provided, as are the number of species 
within each family and the number of those that 
occur in Australia (both native and introduced). 

There are few flaws in the guide. The species 
accounts and illustrations are helpful and well orga- 
nized. It would have been useful, however, to 
include migratory information on the distribution 
maps, perhaps with seasonal ranges depicted with 
separate colours. Distribution maps for some intro- 
duced or rare species are not provided and would 
have been a nice feature. 

As a new resident of Australia, and unfamiliar 
with the birds I was to encounter, I found this field 
guide to be of great assistance. It is small and hardy 
enough to toss into a daypack, but also would make 
a useful reference for anyone interested in Australian 
birds. 

Shawn Morrison 



8/1 Totterdell St, Belconnen, Australian Capital Territory, 
2617, Australia 



Katydids and Bush-Crickets: Reproductive Behavior and Evolution of the Tettigoniidae 



Darryl T. Gwynne. 2001. A volume in the Cornell series in 
arthropod biology (John Alcock, editor). Cornell 
University Press, Ithaca, xii + 317 pp., illus. U.S. $42.50. 

Darwin (1871) introduced the theory of sexual 
selection as an attempt to explain the evolution of 
such traits as bright colours, huge feather plumes, 
antlers, and other male adornments, features that by 
their highly conspicuous nature, seemed highly 
unlikely to confer survival advantages on their bear- 
er. Sexual selection (differential reproductive suc- 
cess resulting from competition for mates) as a topic 
of investigation lay essentially dormant for about a 
century. However, since the mid-1970s, interest in 
sexual selection has blossomed (yes, it occurs in 
plants, too!), following the recognition that it is a 
major evolutionary agent. 

Sexual selection is the underlying theme of 
Gwynne' s book, which in this respect is similar to 
those of Eberhard (1985, 1996), Gould and Gould 
(1989), and Andersson (1994). However, it has the 



advantage over these contributions in that it deals 
with the phenomenon in a specific taxon, the family 
Tettigoniidae, which has allowed the author to pre- 
sent his ideas on a solid platform of basic biology. 
Indeed, the author's successful aim is not only to 
interest students of sexual selection, but to show nat- 
uralists what a fascinating group of insects the tet- 
tigoniids are. 

For field naturalists, the early chapters, which 
deal with evolution, diversity (more than 6000 
species arranged in 17 subfamilies), life histories, 
food preferences, natural enemies, defence mecha- 
nisms, and sound communication, will probably be 
of most interest. Behaviorists, especially those who 
focus on reproductive behaviour, will find much 
food for thought in the second half of the book. 
However, animal biologists in general will find the 
entire volume contains much to hold their attention. 

The book is profusely illustrated with line draw- 
ings, black-and-white photographs (some of which. 



166 



The Canadian Field-Naturalist 



Vol. 116 



unfortunately, lack contrast), and four pages of color 
plates. The text is generally well written and 
Gwynne's approach (present the hypothesis, then 
examine the evidence) is highly recommended as a 
"model" for developing researchers. Occasionally, 
the author "overindulges" by giving too many exam- 
ples to illustrate a point, and the editor might have 
used a sharper pencil to remove redundancy and rep- 
etition and a sharper eye to ensure that the indexing 
is accurate. I stress, however, that these are minor 
"complaints" and my overall opinion is that Gwynne 
deserves full credit for his synthesis of the biology of 
this major, yet not well known, insect group. 



Literature Cited 

Andersson, M. 1994. Sexual Selection. Princeton University Press, 

Princeton, New Jersey. 
Darwin, C. 1871. The descent of man and selection in relation to 

sex. John Murray, London. 
Eberhard, W. G. 1985. Sexual selection and animal genitalia. 

Harvard University Press, Cambridge, Massachusetts. 
Eberhard, W. G. 1996. Female control: Sexual selection by cryptic 

female choice. Princeton University Press, Princeton, New Jersey. 
Gould, J., and C. G. Gould, 1989. Sexual selection. Scientific 

American Library, New York. 

Cedric Gillott 

Department of Biology, University of Saskatchewan, 112 
Science Place, Saskatoon, Saskatchewan S7N 5E2 Canada 



A Manual for Wildlife Radio Tagging 

Robert E. Kenward. 2001. Academic Press, Sai) Diego, 
California. 311 pp., illus. U.S. $65.00. 

This book is a significant updating of ''Wildlife 
radio tagging: equipment, field techniques and data 
analysis'' (Kenward 1987). The book incorporates 
much new information developed since the late 
1980s and includes all of the original topics from the 
earlier volume but is 70% larger. 

Kenward starts this book with the logical least 
common denominator, that is, deciding whether 
radio tagging is an appropriate option for collecting 
data from a subject animal and planning projects. 
Once the decision to radio tag is made, you proceed 
through the chapters of the book to determine the 
appropriate equipment, how to obtain equipment, 
how to make and attach radio tags, proper radio 
tracking technique, and data analysis. 

It is unfortunate that this volume contains many 
errors, particularly in the literature cited. Some of the 
errors can make it difficult for less knowledgeable 
readers to locate and obtain references. 



This book is aimed at the beginning researcher 
contemplating use of radio tagging as well as the 
more experienced. Even with the errors as noted 
above, because of Kenward' s depth of knowledge 
and extensive experience with tracking, this book is 
of definite value to any and all researchers that are 
considering or applying telemetry to their research. It 
is highly recommendable for its thoroughness, time- 
liness, and user friendly organization. Those already 
owning the earlier volume should still consider pur- 
chasing this new version. 

Literature Cited 

Kenward, R. E. 1987. Wildlife radio tagging: equipment, field 

techniques and data analysis. Academic Press, San Diego, 

California. 

Roger D. Applegate 

Research and Survey Office, Kansas Department of 
Wildlife and Parks, P. O. Box 1525, Emporia, Kansas 
66801-1525 USA 



Bird Census Techniques 

By Colin J. Bibby, Neil J. Burgess, David A. Hill, and 
Simon H. Mustoe. 2000. 2nd edition. Academic, 
London. 301 pp., illus. U.S. $55.00. 

The global concerns for the plight of long distance 
migratory birds and efforts to develop comprehen- 
sive conservation plans in North America encom- 
passing all avian species make the need for statisti- 
cally valid bird census techniques evident. This 
book, updating and expanding the first edition, fills 
that need. 

The authors have incorporated the vast literature 
of bird censusing to make this volume a comprehen- 
sive reference. Each major bird counting technique is 



given a thorough treatment in cook-book fashion that 
makes it possible for anyone to lay out and conduct 
the count. The book also provides a general back- 
ground on design of bird censuses and statistical 
issues. There is information on use of computer soft- 
ware provided in the chapter on line transects. 

The book has chapters on applying census meth- 
ods to specific species. These chapters deal with 
species that are largely European. However, in most 
cases, the techniques can be applied to closely relat- 
ed North American species. For example: Willow 
Ptarmigan {Lagopus lagopus) and Red Grouse (L. /. 
scoticus). 



2002 



Book Reviews 



167 



Bibby et al. is relatively free of errors. Those 
noted were minor and do not detract from the read- 
ability or utility of the book. I would like to have 
seen some thoughtful discussion of issues relating to 
the use of index methods as opposed to methods that 
provide an estimate. However, overall this book has 
met the authors' objectives of providing a compre- 
hensive reference on bird censusing. I recommend 
this book to all who are conducting or will be con- 



ducting such field studies. For those owning the first 
edition, I recommend purchasing this second edition. 



Roger D. Applegate 



Research and Survey Office, Kansas Department of 
Wildlife and Parks, P. O. Box 1525, Emporia, KS 66801- 
1525 USA 



Flying Foxes: Fruit and Blossom Bats of Australia 



By L. Hall and G. Richards. 2000. Krieger Publishing 
Company, Malabar, Florida, viii + 133 pp., illus. Cloth 
U.S. $29.50; paper U.S. $21.50. 

I have had a great interest in flying foxes since I 
first saw them roosting in a tree in Indonesia about 
20 years ago. Their size and open nesting sites in an 
urban area made them far more spectacular than the 
Canadian cave species that I had studied as a gradu- 
ate student or even the variety of species I had col- 
lected in the Amazon region of South America. In 
recent years (including the time I am writing this), 
my house in Ibadan, Nigeria, allows me to go out 
every evening and watch spectacular, huge clouds of 
flying foxes migrating overhead from their daytime 
camp in an urban, botanical garden, where they are 
safe from hungry hunters, to their evening fruit-tree 
feeding grounds. 

The flying foxes are fruit and nectar eating bats of 
the suborder Megachiroptera, which are found in the 
old world tropics and are only distantly related to the 
suborder Microchiroptera which I studied as a stu- 
dent. This book did confuse me a bit, at first, since 
the authors primarily discuss only Australian species 
and the most common of these in the genera 
Pteropus, which does not occur in continental 
Africa. The authors state in the introduction "There 
is a number of other fruit-eating Megachiroptera 
which are not in the genus Pteropus but which are 
also called 'flying foxes'. " Then throughout the rest 
of the book they talk of flying foxes as if they were 
only Australian Pteropus. This caused me consider- 
able confusion since I have been involved in some 
flying fox research in Nigeria, and I know this is not 
Australia. Back to my early lessons about how Latin, 
scientific names are so much more precise than com- 
mon names. 

This book is very comprehensive, well re- 
searched, and well written. It provides excellent 
information to those who are interested in flying 
foxes, even if not just in Australia. It covers such 
areas as history and archaeology, identification, 
and distribution of the Australian species, anatomy 



and physiology, disease, behaviour, feeding ecolo- 
gy, migration, conservation, rehabilitation, and 
even hand rearing, in great detail. Appendices 
include native plant foods, extensive references, a 
glossary, and index. There is some confusion 
where references in the text to an individual or 
institution doing research cannot be traced to the 
specific published references at the back of the 
book. 

Some very interesting facts are presented in this 
book. Did you know that bats are the second most 
common mammal group to rodents. Then there is the 
blossom bat, described as a "mouse-sized flying fox" 
— a bit of a contradiction. Flying foxes, unlike other 
bats, rely largely on sight. They will not fly in total 
darkness and do not hear much better than a human 
being. Their vision is similar to that of a cat. During 
the day, flying foxes roost together in large "camps" 
which can number over 250 000. These camps can 
be very noisy and result in damages to the local 
trees. At night, they will migrate long distances to 
feed, again causing damages due to their numbers. 
This has resulted in cropping by orchard owners, 
which has decimated the population in Australia. 
They are now protected and conserved for their ben- 
eficial behaviours such as pollination and dispersal 
of fruit seeds. 

Flying foxes have learned the behaviour of roost- 
ing in urban forests or parks as a protection against 
shooting. In my area of Nigeria, the largest local 
camps are the botanical garden close to our house, 
and two local university campuses. Flying foxes are 
considered a culinary delicacy in Nigeria as well as 
elsewhere in their range. One of my co-workers here 
says they make delicious soup. But he was surprised 
that they do not taste like a bird. Usually if I eat bush 
meat, I do not ask what it is. 

With a lifespan (in captivity) of up to 50 years, 
flying foxes only produce one young per year. This 
has resulted in serious population drops due to 
habitat loss and hunting pressure. Several southeast 
Asian islands have seen populations become extinct 



168 



The Canadian Field-Naturalist 



Vol. 116 



and the entire population of some species in 
Australia is now smaller than the 300 000 reported 
in one camp less than fifty years ago. Another 
species which 1 hope will not disappear before we 
have really had the chance to get to know it. This 
book is a very worthwhile and authoritative source 



of information on the intriguing world of flying 
foxes. 

Wilson Eedy 

Geomatics Nigeria Limited, Plot 7 Off DPC Road, Ibadan, 
Nigeria 



Botany 

Guide to Standard Floras of the World: An Annotated Geographically Arranged Systematic 
Bibliography of the Principal Floras, Enumerations, Checklists, and Chorological Atlases of 
Different Areas. Second Edition. 



By David G. Fordin. 2001. Cambridge University Press, 
The Edinburgh Building, Cambridge CB2 2RU UK; 40 
West 20th Street, New York, NY 10011-4211, U.S.A. 
xxiv + 1 100 pages U.S. $240.00 + $6.00 postage. 

This volume, which includes information on all 
the floras of the world known to the author up to the 
end of the twentieth century, is a tremendous step 
ahead of the first edition which was published in 
1984 and numbered 619 pages. The first pages of 
this volume include prologues and acknowledge- 
ments of the first and second editions; Part I which 
numbers 84 pages and includes an analytical- 
synthetic systematic bibliography of "standard" flo- 
ras: scope, sources and structure, the evolution of 
floras, floras at the end of the twentieth century: phi- 
losophy, progress, and prospects and references. Part 
II includes systematic bibliography, conspectus of 
divisions and superregions, world floras, isolated 
oceanic islands and polar regions, and divisions: 
North America (north of Mexico), Middle America, 
South America, Australia and islands of the south- 
west Indian Ocean (Malagassia), Africa, Europe, 
northern, central and southwestern (extra-monsoon- 
al) Asia, southern, eastern and southeastern (mon- 
soonal) Asia, Greater Malasia, and Oceania. This is 
followed by Appendix A: major general bibliogra- 
phies, indices, and library catalogues covering world 
floristic literature, and Appendix B: Abbreviations of 
serials cited, addenda in proof, geographical index, 
and author index. 

The main part of this volume then proceeds through 
the areas listed in Part II above with descriptions of 
the regions, locations, number of species, early botani- 
cal history up to the end of the twentieth century, and 
the major floras which have been published through 
the years. There is a wealth of information here both 
for those living in any region or anyone interested in 
the botany of another part of the world. 

It is inevitable in a work such as this that occa- 
sionally some useful publications will be passed by. 
Some from northern North America that might have 
been included are the following: 



Benson, L. 1982. The Cacti of the United States and 

Canada. Stanford University Press. 
Biek, D. 1999. Flora of Mount Rainier National Park. 

Oregon State University Press. 
Cody, W. J. 1988. Plants of Riding Mountain National 

Park, Manitoba. Agriculture Canada. 
Douglas, G. W., C. B. Straley, and D. V. Meidinger. 

1998. Rare Native Vascular Plants of British Columbia. 

Crown Publications. 
Hallworth, B., and C. C. Chinnappa. 1997. Plants of 

Kananaskis Country in the Rocky Mountains of Alberta. 

University of Calgary Press. 
Holmgren, A. H. 1998. "Illustrated Companion to 

Gleason and Cronquists' Manual: Illustrations of the 

Vascular Plants of Northeastern United States and 

Adjacent Canada". The New York Botanical Garden. 
McGregor, R. L., and T. M. Barkley. 1977. Adas of the 

Flora of the Great Plains. The Iowa State University 

Press. 
Packer, J., and C. E. Bradley. 1984. Checklist of the rare 

plants in Alberta. Provincial Museum of Alberta 

National History Occasional Paper Number 5. 
Petrik-Ott, A. J. 1979. The Pteridophytes of Kansas, 

Nebraska, South Dakota and North Dakota. Nova 

Hedwigia, Beiheft 61. 
Argus, George [Series Editor] 1977-1995. Rare Native 

Vascular Plants of Canadian Provinces and Territories. 

Syllogeus National Museum of Natural Sciences, 

Ottawa. 
Tryon, A. F., and R. C. Moran. 1997. The Ferns and 

Allied Plants of New England. Centre for Biological 

Conservation, Massachusetts Audubon Society. 

Botanical studies will continue around the world 
and already one for the year 2000 has been published 
in Canada: H. R. Hinds, Flora of New Brunswick, 
Second Edition, which was a great step ahead from 
the First Edition. It will be most important to keep 
The Guide to Standard Floras of the World up to 
date through new editions or supplements. 

William J. Cody 



Eastern Cereal and Oilseed Research Centre, Agriculture 
and Agri-Food Canada, Wm. Saunders Building, Central 
Experimental Farm, Ottawa, Ontario KIA 0C6 Canada. 



2002 



Book Reviews 



169 



EVIRONMENT 

Their Fathers' Work: Casting Nets with the World's Fishermen 



By W. McCloskey. 2000. International Marine/McGraw- 
Hill, New York. 370 pp. U.S. $20.95 

This book provides the reader with a superb and 
highly praised overview of global fisheries, focusing 
on Alaskan waters. In addition, it also covers first- 
hand experiences for offshore and coastal fisheries 
with vessels from Japan, Chile, Indonesia, New- 
foundland (Grand Banks), Maine (Georges Bank), 
Iceland, and Norway. The book is very pleasant to 
read as it combines fiction with fact. It is a heroic and 
romantic description of a likely soon-to-be-gone life 
of hard work. Nevertheless, reading how other people 
work very hard and under life-threatening conditions 
might present some sort of decadence; but so be it. 

In case the reader is not familiar with how to 
cheat in this business of international fisheries and 
quotas (led by Spain, Taiwan, Japan, and many East 
European nations) this book will be enlightening. It 
outlines in detail how fishing quotas are easily dou- 
bled, if not ignored, by many vessel captains and 
fishermen worldwide. The explicit use of Dynamite 
Fishing, Liner Nets (an additional net with an ille- 
gally smaller mesh-size put inside the regular net), 
the "Pareja" Method (one huge net pulled by two 
boats), and many other tricks are shown and suggest- 
ed: e.g., the same vessel being registered with two 
different names (thus, multiplying the quota by two), 
stowing an additional catch somewhere under deck, 
trading the catch offshore (therefore enabled to start 
again with a "new" quota), and mis-reporting catch- 
es. When fisheries officers appear for control and 
gear inspection, nets simply get cut off (which 
makes it even worse for fish, seabirds, and sea mam- 
mals that drown in the "ghost nets" later). Overall, I 
find that the author, an American, might have a ten- 
dency to put too much blame on the Spaniards and 
Russians. Instead, a mention and description of the 
role that the Vladivostok-based Russian fisheries 
play, acting worldwide, could have made the book 
even better. 

The thorough understatement of environmental 
damage done by coastal and offshore fishery must 
be of concern to any informed naturalist. The author 
neglects to address the destructive fishery method 
from draggers ("seafloor dredging"), which is esti- 
mated to damage an area larger than that lost 
through deforestation in the tropics. There is no 
mention of fisheries gear polluting beaches world- 
wide, or "ghost nets" which float around the 
world's oceans for years (eventually, they will sink, 
but only the fish know whether they will ever rot). 
Sensitive by-catch topics such as the endangered 
Short-tailed Albatross {Phoehastria albatrus) 
caught by freezer-longliners fishing off Alaska are 
not mentioned, and certainly there is no reporting of 



the numerous sea turtles, sharks, dolphins, porpois- 
es, seabirds, moon fishes, and many other species 
suffering and dying for the sake of high quality fish. 
In times of environmentalism, that might be seen as 
a short-coming of this book. Although the occur- 
rence of a "black catch" is somewhat mentioned, 
one reads that shrimp fisheries have apparently 
almost no by-catch. The reader has to hold his/her 
breath when McCloskey mentions "overpopula- 
tions" of Sockeye and seals; 50 000 seals are 
described as an "overpopulation" rather than vic- 
tims in a potential by-catch problem. No wonder, 
the author identifies clearly from the "fishermen's 
side", blames Greenpeace, and does not place fish- 
eries in the overall context of the environment. 
Instead, he largely focuses on economical and 
descriptive aspects of fisheries. In this regard, the 
author's presentation of Chile's fishery develop- 
ment lacks sensitivity to the well-proven and nega- 
tive effects of over-commercialization. On the other 
side, his wonderful and detailed presentation of the 
effects from the Exxon Valdez Oilspill for Alaska 
and its island communities compensate for the pre- 
vious short-comings. A remarkable link is shown 
why the prizes of the Japanese salmon market are 
driven by cycles of the Japanese salmon runs and 
thus dictate the Alaskan salmon fisheries. 
McCloskey gets closer to the heart of the fisheries 
problem when outlining that improved efficiency 
and introduction of very light, and therefore allow- 
ing for longer, plastic nets has contributed to the 
current overfishing crisis. 

In the numerous and fascinating chapters the author 
also emphasizes and describes that there exists such a 
thing as severe overfishing: Snow Crab in Alaska; 
Cod, Flounder and Squid in Newfoundland; and 
Halibut off West America. He blames governmental 
mis-management and elaborates nicely throughout the 
text that there is also conflict of interest among fisher- 
men on these topics; e.g., unions, and small scale fish- 
eries vs. industrialized trawlers. In the context of gov- 
ernmental mismanagement, New Zealand's Orange 
Roughy, a prime example of overfishing and disas- 
trous fisheries management, could have been men- 
tioned, too. The book would have been even better if 
topics such as a Native Fishery Rights, North Sea 
Fisheries, and Krill Fisheries in the Antiyctic had been 
included. The map of the Grand Banks lacks the 
French Fisheries zone around St. Pierre and 
Miquelon; but the reader will appreciate that this book 
has a very detailed index, which allows that it can 
serve as a valid source of references. 

The book ends with a well-written and conclu- 
sive section on global fisheries and policy. The 
author quotes from one of his many interviews with 



170 



The Canadian Field-Naturalist 



Vol. 



experts: ''Gathering fishery statistics is an art in 
probability". That statement makes it clear that, cur- 
rently, there can be no sustainable world fisheries. 
Due to the many topics covered, 1 thoroughly 
enjoyed reading this book and got literally "hooked". 



Falk Huettmann 

Centre for Wildlife Ecology, Biology Department, Simon 
Eraser University, 8888 University Drive, Bumaby, British 
Columbia V5A 1S6 Canada 



AAAS Atlas of Population and Environment 

By P. Harrison and F. Pearce. 2001. University of 
California Press, Berkeley, xi + 204 pp., illus. Cloth U.S. 
$65; paper U.S. $29.95. 

Population is the main cause of environmental 
problems. This book provides an excellent, well- 
documented, and authoritative demonstration of this 
principle, using easy to read text and great atlas dis- 
plays. It needs to be read by every person who is 
concerned over our world environmental degradation 
and the population impacts of the very near future. It 
comes home to me personally, spending much time 
in Nigeria, one of the highest and most densely pop- 
ulated countries in Africa. In spite of the HIV-AIDS 
epidemic, it still has one of the highest population 
growth rates in the world (2.81% per year). Here one 
comes face to face with poverty, lack of clean water 
supplies, major losses of forest and ecological habi- 
tat, desertification, and the desperation of many peo- 
ple struggling to survive from day to day. I see 
things that I never would have believed before I first 
came here 10 years ago. What will it be like in 2050 
AD when the population is expected to grow by 
more than 50% to about 200 million people (in an 
area the size of Ontario, with large areas of arid 
savanna). This book gives some predictive outlook 
for such areas of the world, where the major popula- 
tion growth will be in the future. I do not think this is 
an issue that we can afford to ignore. Perhaps, as a 
bit of a field-naturalist heretic, I suggest that it is 
more of an environmental priority than the birds in 
our back yard. I recommend this book as a critical 
reference, resource for anyone concerned about the 
environment and the future of our world. 

The book is organized into three major sections. 
Part 1 : Overview (40 pages) is primarily introducto- 
ry text, covering issues such as environmental poli- 
cies and philosophies, the scale of our existence, and 
the theory of population-environment links. How 
many people can the world sustain? This area is not 
well understood and the answer is not given, but the- 
ories such as those of Malthus and Erhlich are ana- 
lyzed in an impartial scientific manner. Much data is 
presented, but the reader is often left to make their 



own conclusions. In the last century population 
expansion has occurred at an alarming, unprecedent- 
ed rate (from three million — 2000 years ago, to 1 
billion 200 years ago, to over 6 billion now, and pre- 
dictions to over 9 billion by 2050). In the last centu- 
ry, we have lost/of the world's topsoil, 1/5 of our 
agricultural land base, major parts of our forest, and 
other habitat. We have experienced a species extinc- 
tion rate 100 times that of any time in our previous 
history. Our problems are summarized in a rather 
sickening and scary manner, but easily understood 
by those of us who live much of our lives in the third 
world. 

The second part of the book is 164 pages of an 
atlas of resources and resource uses or losses, along 
with some accompanying discussion and references. 
Areas covered include population effects on natural 
resources, land use, atmosphere, wastes and chemi- 
cals, ecosystems, and biodiversity. The atlas maps 
allow easy reference and the text provides excellent 
detail. 

The third part of the book provides a few case his- 
tories of population-environment interaction, for dif- 
ferent biogeographic regions of the world. All but 
one are taken from World Wildlife Fund informa- 
tion. The one exception, and location closest to 
Canada, is a Nature Conservancy discussion of the 
Sonoran Desert (a favourite retreat for me). 

I guess Canada can be happy to be on the good 
side of most of these environmental issues and to be 
proud not to have a case history illustrated here. 
However, I do strongly believe that we cannot hide 
from these potential future population-caused envi- 
ronmental catastrophes. We are all in one world and 
the information in this book is an excellent start to 
planning to save it. 

Wilson Eedy 

Terfa Inc, RR# 1, Glencaim, Ontario LOM IKO Canada and 
Geomatics Nigeria Limited, Plot 7 Off DPC Road, 
Ibadan, Oyo State, Nigeria (terfa@geoniger.com, 
www.geoniger.com) 



2002 



Book Reviews 



171 



Making Better Environmental Decisions 

By Mary O'Brien. 2000. The MIT Press, Cambridge, 
Massachusetts. 286 pp., illus. U.S. $22.95. 

This book starts with a story, a parable of sorts. A 
woman is standing by an icy mountain river, intend- 
ing to cross to the other side. A team of risk asses- 
sors accompanies her, reviewing her situation. They 
present arguments and calculations, concluding that 
the risk she faces is low from their various perspec- 
tives — toxicology, cardiology, hydrology, policy — 
and that she should wade across the river. Yet, to 
their extreme frustration, she refuses to wade. Why? 
She points upstream and tells them, "Because there 
is a bridge." 

The risk assessors evaluated the risk of only one 
option: wading across the river. The woman 
reviewed her alternatives, which included crossing 
on the bridge upstream. Making Better Environ- 
mental Decisions is about doing what the woman in 
the parable did: making decisions based on all possi- 
ble alternatives instead of considering only the risk 
associated with a narrow range of options. 

The concept of alternatives assessment is based on 
certain common-sense principles. These include, 
among others, that it is unacceptable to harm human 



or non-human life when reasonable alternatives 
exist. That nobody should be able to define for any- 
one else how much damage is "acceptable". That 
those affected by decisions should be involved in 
making them. And that there are always alternatives 
that can be considered. 

The book outhnes why risk assessment — the pro- 
cess currently relied on by governments and the pri- 
vate sector to make decisions — doesn't work, and 
goes on to explain why alternatives assessment is bet- 
ter and how to make the shift. Clearly written and 
easy to read, the book is full of convincing examples, 
including many relating to ecosystems and wildlife, 
that speak loudly for considering all possible alterna- 
tives before making decisions that will affect the 
health of humans and the natural environment. 

This book is a must-read for all naturalists 
involved in decisions-making processes, as profes- 
sionals or as volunteers, and for anyone concerned 
about the future of life on this planet. 



R. Sander-Regier 



RR5 Shawville, Quebec JOX 2Y0 Canada 



Scientific Method for Ecological Research 

By E. David Ford. 2000. Cambridge University Press, 
Cambridge. 564 pp., illus. Cloth U.S. $10; paper U.S. 
$49.95 

Dr. Ford, professor, College of Forest Resources, 
University of Washington, carves out, in exemplary 
fashion, a philosophy of ecological research. The 
philosophical exploration of the "hard" physical sci- 
ences as the paradigmatic sciences introduced 
assumptions as to what makes "good science," but it 
has become evident that certain of these assumptions 
are questionable or even false when it comes to the 
biological sciences and ecology. The philosophy of 
the physical sciences has created a "storybook scien- 
tist" who is considered specialized, reductionist, sci- 
entifically objective, rigorous, experimental, predic- 
tive, and essentially value-free. For Professor Ford, 
such a description is a caricature. Ford eschews any 
scientific method which defies particular techniques, a 
particular form of scientific reasoning, asserts that cer- 
tain questions are more worthwhile than others, or 
assumes that questions must be asked in specific 
ways. For the author, the scientific method is not an 
immutable, monolithic process characterized by true 
and tried roadsides, but a creative process in which 

I these roadsigns may take many and varied forms 
depending on the particular research question at hand. 
Written with the graduate student and aspiring sci- 



into four main sections; an analytical framework for 
conducting research, the basis for scientific inference 
and theory development in ecology, the social 
dimensions of research, and finally the presentation 
of a methodology for developing ecological theory, 
his so-called progressive synthesis. Making explicit 
how ecologists think in their research is his over- 
arching goal. Of significance to Ford is a recognition 
of the unique features of ecological research which 
demand a research methodology different from that 
assumed for the physical sciences. Of particular con- 
cern is the open or partially open nature of ecologi- 
cal systems, the probabilistic nature of many ecolog- 
ical concepts, the inherent variability of ecological 
data, and problems of multiple causality. All of these 
factors pose greater difficulties in measurement and 
concept development, conditions for scientific infer- 
ence, and the basis for theory acceptance or refu- 
tation when compared to controlled, laboratory 
conditions common to the physical sciences. 

Ford's pedagogically refined review of the 
diverse forms of scientific reasoning and philosoph- 
ical assumptions grounding scientific research is 
exhaustive and brimming with insight. Lack of 
space prevents the consideration of many critical 
issues, but attention to a select few is profitable. 
Particular emphasis is placed by Ford on the critical 
analysis of concepts and propositions, both of which 



172 



The Canadian Field-Naturalist 



Vol. 116 



will act as the foundation of research. What is 
known (axioms), what one wishes to know (postu- 
lates), and how one intends to collect data that will 
assess postulates (data statements) are critical first 
steps in research. The clarification of ecological 
concepts is important as well. Ecology often focuses 
on what Ford terms integrative concepts (e.g., sta- 
bility, ecological integrity, resilience), theoretical 
constructs about ecological organization that defy 
direct measurement and demand synthesis from a 
variety of system studies. Caution is directed to the 
uncritical use of statistical inference as the sole 
assessment of postulates. Highly significant is the 
notion of theory domains which define the bound- 
aries within which a particular theory may or may 
not operate, effectively rendering ecological theory 
development very complex. Of significance as well 
is the notion that both measurement and experiment 
are essentially an art. Even when precise and accu- 
rate, all measurements by definition are abstracts in 
that they represent the object of interest, but are not 
the things themselves. Ford refutes the notion that 
Popperian falsification can be used as the basis for 
theory change, citing evidence to show that the 
rational basis for theory acceptance or rejection is 
much more complex. 

The reader is introduced to the "subjective" or 
sociological dimensions of science. Science is pre- 
sented as not simply an automated, invariant or 
determined process, but as a human activity under- 



taken by men and women acting as bearers of wis- 
dom, passion, faults and foibles seemingly no more 
or no less than the "average citizen." Ford addresses 
such issues as scientific fraud, the publication pro- 
cess, peer-review, the significance of gender, and the 
role of scientists in developing public policy. Social 
processes are viewed as significant in determining 
which questions or research problems are considered 
important, which group will receive funding 
resources and precisely how research questions are 
formulated and pursued. 

A popular view of science may assume that the 
scientific enterprise is a wholly "objective" process 
to which the particular investigator simply submits. 
Professor Ford returns a human face to science 
emphasizing the role of creativity, choice, probabili- 
ties, the art of science, plurality of logical reasoning 
and methodological ideals, a distancing from a priori 
philosophical assumptions. This by no means weak- 
ens the scientific enterprise, but rather re-introduces 
a rigorous and critical attitude in posing scientific 
problems and developing an adequate procedure of 
assessment. In doing this, Professor Ford ennobles 
human scientific activity by calling explicit attention 
to the pattern of scientific reasoning. 

John McCarthy, S.J. 



St. Mark's College, University of British Columbia, 5935 
lona Drive, Vancouver, British Columbia V6T 1J7 Canada 



Miscellaneous 

The Last Great Sea: 

A Voyage through the Human and Natural History of the North Pacific Ocean 



By T. Glavin. 2000. David Suzuki Foundation and 
Greystone Books, Douglas & Mclntyre Publishing 
Group, Vancouver/Toronto. 244 pp. 

This book belongs in every conservationist's 
bookshelf, to say the least. As D. Suzuki describes 
very convincingly in the foreword, the environment 
of today's North Pacific is characterized by its loss 
of (fish) species and its wipe-out of protein assem- 
blages. The collapse of Sockeye Salmon is only one 
of many sad examples, many more exist: Steller's 
Sea Cow, Spectacled Cormorant, Dwason's Caribou 
(Queen Charlotte Islands), and even plant species 
like Tobacco (Queen Charlotte Islands). Other 
species like Walruses, Sea Otters, and Fur Seals have 
barely survived until now. 

The first chapter starts slowly but allows a very 
solid overview about historical and archaeological 
facts. After Chapter 2, nobody can deny anymore the 
environmental disaster and mis-management of the 



North Pacific and coastal British Columbia. 
Nevertheless, the author convinces the reader that the 
North Pacific still is THE largest fish producer in the 
world. ''As in aboriginal fisheries, mythology played a 
part in industrial fisheries management, especially the 
myth of a superabundant ocean and the all-powerful 
capability of science and technology to fix the messes 
made by hydroelectric dams, lousy forestry practices 
and overfishing"". The governmentally encouraged 
Merganser Control and Bear Shooting Programs 
designed for the sake of Salmon Protection prove this 
citation very well. Galvin strongly eliminates all illu- 
sions on how to heal the problem of overfishing. For 
instance, he shows that S. Livingstone's widely fol- 
lowed idea of Fish Hatcheries does not produce more 
salmon, but instead takes away funds and harms natu- 
ral salmon stocks since they simply replace the last 
remaining and struggling stocks with poorly adjusted 
new ones; Gavin's arguments are also strong against 



I 



2002 



Book Reviews 



173 



Salmon Farming; e.g., it contributes to the closure of 
marine fisheries for wild salmon, and it requires 3 kg 
of fish to produce 1 kg of salmon. 

All Pacific salmon species are discussed: Chum, 
Sockeye, Pink, Coho, Steelhead, Masu and Amago. 
Of major interest is in this regard the scientific dis- 
cussion around the taxonomy of salmon; e.g., 
Steelhead (classified until 1980s as Trout). The 
author brilliantly points out the implications of the 
religious-based and somewhat outdated taxonomical 
system by Carl von Linne, and how this affects the 
species management by national governments 
(provincial and federal) on an international level 
even (Canada vs. U.S. A). 

The backwardness and failure of fishery laws are 
shown by outlining that the first salmon-fishing reg- 
ulations for the Fraser River was a simple word-for- 
word replication of fishing regulations on English 
Rivers. At that time, Canada's external affairs juris- 
diction was still controlled by the British, which 
affects the Canada-U.S. salmon treaty concluded 
1930s and renewed in 1985. In addition, Galvin 
shows that Canadian and U.S. fishery scientists sig- 
nificantly differed in their stock assessment results 
for the same species in the same waters even; conse- 
quently, so did the management and political agen- 
das. This is the classical picture of "mixed-stock" 
fisheries, which also threatens small salmon runs. 

The author reports the incidental death of 50 000 
marine mammals and 500 000 seabirds due to driftnet 
fishery activities in the North Pacific; marine (plastic) 
pollution comes with it. Despite the well shown fail- 
ure of a European and Western approach dealing with 
the North Pacific fisheries, domestic Japanese, and 
Native fisheries seemed to work well and sustainable. 
Galvin shows the magnitude of "pre-contact" fishery 
for salmon by natives, which was even comparable 
with levels of commercial fisheries from this century. 
Some readers might find that the book slightly fol- 
lows stereotypical views of the noble native. 

A very strong point in this book is how the North 
Pacific and its fauna is linked with the "hinterland": 
Old-growth rainforest, landscape and Bald Eagles. 
This needs to be considered in the light that resident 
Killerwhales in British Columbia are among the 
most contaminated cetaceans of the world. 

A very complete picture of the North Pacific is 
portrayed by fully considering the Russian influence 
and history. The book outlines well that Russian set- 
tlers did much better than the western type of colo- 
nization (a point that might be put in doubt for the 
Kodiak Islands at least). The Russian-American 
Company was much more relevant in the history of 
North Pacific settlements and explorations than the 



Hudson Bay Company (HBC). But nevertheless, as 
with the HBC, the Russian quest for the North 
Pacific had the same motivation: central European 
pelt resources were already overhunted! 

Regarding the marine ecology of the North Pacific, 
the importance of the Aleutian low. Pacific currents, 
and El Nino are fully described. This ecosystem is 
driven by 'regime changes', which calls for a dynam- 
ic management. The author outhnes this very well by 
presenting the ground-breaking work from Russian 
Scientist T. Baranov, but also from Bill Ricker 
("Ricker curve") and others at the Pacific Biological 
Station; e.g., G. McFarlane and D. Beamish. A quote 
from the book says it all: Understanding catch statis- 
tics is like ''reading a single faded and crumbling 
onionskin page from an early draft of Wagner's 
Tannhaeuser, in a dimly lit room'\ Another quote of 
the book and taken from the U.N. Code of Conduct 
for Responsible Fisheries states, in part, that ''the 
absence of adequate scientific information should not 
be used as a reason for postponing or failing to take 
conservation and management measures'". Well said. 

The chapters on anthropology and human history 
of the North Pacific and how the Russians, Asians, 
and Natives settled and explored the North Pacific 
are on the same level than high-caliber books as 
Guns, Germs and Steel by J. Diamond. Just to name 
some highlights, Glavin mentions how natives grew 
Arrowhead and potatoes, he cites the work of the 
Russian Anthropologist S. Fedorova, and he docu- 
ments that Hawaiians, Japanese, Chinese and 
Russians presented a major group of settlers. In addi- 
tion, the book reports a lot of British Columbian and 
Vancouver history and puts Canada in the context of 
the overall Pacific. 

Despite the fact that whahng, sealing, and eating 
dolphins are as old as the human history of the North 
Pacific, whale watching (starting as early as 1907) 
has already produced more profit than commercial 
whaling ever did for western North America. Green- 
peace started in Vancouver; it "was bom in the blood 
of whales". Nevertheless, the author shows that 
already in the 19th century the pelagic seal hunt pro- 
voked the first great international controversy about 
the overharvesting of the world's marine mammals. 
It resulted in the international milestone contract 
('fur seal treaty') of 1911 between Russia, Japan. 
Canada and U.S.A. 

Falk Huettmann 

Centre for Wildlife Ecology, Biology Department, Simon 
Fraser University, 8888 University Drive, Bumaby. British 
Columbia V5 A 1S6 Canada 



174 



The Canadian Field-Naturalist 



Vol. 116 



New Titles 



Zoology 



Animal ecology. 2001. By C. Elton. University of 
Chicago Press, Chicago, xvii + 209 pp., illus. U.S. $20. 

*A birder's guide to metropolitan areas of North Amer- 
ica. 2002. Edited by P. Lehman. American Birding 
Association, Colorado Springs, iv + 508 pp., illus. 

tThe birds of heaven: travels with cranes. 2001. By P. 
Matthiessen. Greystone Books, Douglas and Mclntyre, 
Vancouver, xv + 349 pp., illus. $36.95. 

*The complete guide to the birds of Europe. 2000. By 
K. Mullarney, L. Svsnsson, D. Zetterstrom, and P. J. 
Grant. Princeton University Press, Princeton. 399 pp., 
illus. U.S.$49.50. 

t Conversations with an eagle: the story of a remarkable 
relationship. 2002. By B. Cox. Greystone Books 
(Douglas and Mclntyre), Vancouver. 288 pp. $22.95. 

tCoral reef fishes. 2002. By E. Lieske and R. Myers. 
Revised edition. Princeton University Press, Princeton. 
400 pp., illus. U.S.$24.95. 

*The Cuban treefrog in Florida. 2001. By W. E. Meshaka, 
Jr. University Press of Florida (Canadian distributor 
Scholarly Book Services, Toronto), xxiii + 191 pp., illus. 

*Dinosaurs: the encyclopedia, supplement 2. 2002. By 
D. F. Glut. McFarland, Jefferson, North Carolina, x + 685 
pp., illus. U.S.$75. 

The family butterfly book: projects, activities, and a 
field guide to 40 favorite North American species. 

2001. By R. Mikula. Storey Books, Pownal, Vermont, x + 
166 pp., illus. Cloth U.S.$29.95; paper U.S.$16.95. 

tGeographic variation in size and shape of savannah 
sparrows (Passerculus sandwichensis). 2001. By J. D. 
Rising. Cooper Ornithological Society, Camarillo, Cali- 
fornia. 65 pp., illus. U.S. $7. 

The grizzly almanac. 2001. By R. H. Busch. Lyon 
Press, New York. 240 pp., illus. U.S.$29.95. 

*Guia de las aves de Espana. 2000. By E. de Juana and 
J. M. Varela. Lynx Edicions, Barcelona. 223 pp., illus. 15 
Euros. 

*Handbook of the birds of the world: volume 7: jaca- 
mars to woodpeckers. 2002. Edited by J. de Hoyo, A. 
Elliot, and J. Sargatal. Lynx Edicions, Barcelona. 613 pp., 
illus. U.S.$185. 

*National Audubon Society guide to marine mammals 
of the world. 2002. By R. R. Reeves, B. S. Stewart, P. J. 
Clapham, and J. A. Powell. Alfred A. Knopf, New York. 
527 pp., illus. U.S. $26.95. Canadian distributor Random 
House, Toronto. $39.95. 

Neotropical treeboas: natural history of the Corallu hor- 
tulanus complex. 2002. By R. W. Henderson. Kreiger 
Publishing, Melbourne, Florida. 228 pp., illus. U.S.$44.50. 



New animal discoveries. 2001. By R. Orenstein. Twenty- 
First Century Books, New York. 64 pp., illus. U.S.$25. 

Noises in the night: the habits of bats. 2001. By D. 
Kovacs. Steck- Vaughn, Austin, Texas. 48 pp., illus. 
U.S. $25. 

Pheasants, partridges, and grouse. 2002. By S. Madge 
and P. McGowan. Princeton University Press, Princeton. 
480 pp., illus. U.S. $49.50. 

A pocket guide to reptiles and amphibians of Alberta. 

2002. By A. P. Russell and A. M. Bauer. Red Deer Press, 
Calgary. 260 pp., illus. $24.95. 

Sea soup: zooplankton. 2001. By M. M. Cerullo. Til- 
bury House, Gardiner, Maine. 40 pp., illus. U.S. $16.95. 

tSharks. 2002. By A. and A. Ferrari. Firefly Books, 
Willowdale, Ontario. 256 pp., illus. $24.95. 

Tadpoles of south-eastern Australia: a guide with keys. 

2002. By M. Austis. Kreiger Publishing, Melbourne, 
Florida. 281 pp., U.S.$44.50. 

*Tales from the underground: a natural history of sub- 
terranean life. 2001. By D. W. Wolfe. Perseus, Cam- 
bridge, Massachusetts, x + 223 pp., illus. U.S. $26. 

tThunder on the tundra: Inuit qaujimajatugangit of the 
Bathurst caribou. 2002. By N. Thorpe, N. Hakongak, S. 
Eyegetok, and the Kitikmeot Elders. Available through 
Natasha Thorpe, 231 Irving Road, Victoria, B.C. V8S 
4A1.208pp., illus. $42. 

tTurtles and tortoises. 2002. By V. Ferri. Firefly Books, 
Willowdale, Ontario. 256 pp., illus. $24.95. 

Why elephants have big ears: understanding patterns 
of life on earth. 2001. By C. Lavers. St. Martins, New 
York. 288 pp., illus. U.S.$23.95. 

Botany 

*Dlctionary of the fungi. 2001. By P. M. Kirk, P. F. 
Cannon, J. C. David, and J. A. Staples. CAB International 
(distributed by Oxford University Press, Cary, North 
Carolina), xi + 655 pp., illus. 

A field guide to water and wetland plants of the 
prairies. 2002. By H. Lahring. Red Deer Press, Calgary. 
192 pp., illus. $18.95. 

tForest dynamics and disturbance regimes: studies 
from temperate evergreen — deciduous forests. 2002. 
By L. E. Frelich. Cambridge University Press, New York, 
ix + 266 pp., illus. U.S.$70. 

Harmful algal blooms on the North American west 
coast. 2001. Edited by R. Ralonde. Alaska Sea Giant 
College Program, Fairbanks. 73 pp. U.S$10. 



2002 



Book Reviews 



175 



tNew world botany. 2001. By R. H. Peterson. Koeltz 
Scientific Books, Koenigstein, Germany. xv+ 638 pp., 
illus. Cloth Euro 93; paper Euro 62. 

Wildflowers around the year. 2001. By H. Ryden. 
Clarion, New York. 90 pp., illus. U.S.$17. 

Wild flowers of field and slope. 2002. By L. Clark. 
Harbour Publishing, Madeira Park, British Columbia. 
80 pp., illus. $9.95. 

Environment 

The B.C. roadside naturalist. 2002. By R. Cannings 
and S. Cannings. Greystone Books (Douglas and 
Mclntyre, Vancouver). 240 pp., illus. $29.95. 

The beachcombers's guide to seashore life of Cali- 
fornia. 2002. By J. D. Sept. Harbour Publishing, 
Madeira Park, British Columbia. 300 pp., illus. CAN 
$28.95; U.S.$17.95 

t Conserving living natural resources in the context of a 
changing world. 2002. By B. J. Weddell. Cambridge 
University Press, New York, xvi + 426 pp., illus. U.S. $35. 

Evolution's workshop: God and science on the 
Galapagos Islands. 2001. By E. J. Larson. Basic Books, 
New York. 322 pp., illus. U.S.$27.50. 

The functional consequences of biodiversity: empirical 
progress and theoretical extensions. 2002. Edited by 
A. P. Kinzig, S. W. Pacola, and D. Tilman. Princeton 
University Press, Princeton. 392 pp., illus. Cloth U.S.$75: 
paper U.S.$29.95. 

Green phoenix: restoring the tropical forest of 
Guanacaste, Costa Rica. 2001. By W. Allen. Oxford, 
New York. 310 pp. U.S.$35 

tintroduction to conservation genetics. 2002. By R. 
Frankham, J. D. Ballou. and D. A. Briscoe. Cambridge 
University Press, New York, xi + 617 pp., illus. Cloth 
U.S. $130; paper U.S.$50. 

tLife at the limits: organisms in extreme environments. 

2002. By D. A. Wharton. Cambridge University Press, 
New York, xi + 307 pp., illus. U.S.$25. 

*Making parks work: strategies for preserving tropical 
nature. 2002. Edited by J. Terborgh, C. van Schaick, L. 
Davenport, and M. Rao. Island Press, Washington, xix + 
511 pp., illus. Cloth U.S.$65; paper U.S.$32.50. 

Marine community ecology. 2001. Edited by M. D. 
Bertness, S. D. Gaines, and M. E. Hay. Sinauer, Sunder- 
land, Massachusetts. 550 pp., illus. U.S. $59.95. 

tMultitrophic level interactions. 2002. Edited by T. 
Tschrntke and B. A. Hawkins. Cambridge University 
Press, New York, vii+274 pp., illus. U.S.$75. 

*Politics of the wild: Canada and endangered species. 

2001. Edited by K. Beazley and R. Boardman. Oxford 



University Press Canada, Toronto, x + 254 pp., illus. 

$27.95. 

tReef life. 2002. By A. and A. Ferrari. Firefly Books, 
Willowdale, Ontario. 288 pp., illus. $24.95. 

tSpread sheet exercises in conservation biology and 
landscape ecology. 2002. By T. M. Donovan and C. W. 
Welden. Sinauer Associates, Sunderland, Massachusetts. 
xi + 464pp., illus. U.S.$5. 

tSpread sheet exercises in ecology and evolution. 2002. 
By T. M. Donovan and C. W. Welden. Sinaeur Associates, 
Sunderland, Massachusetts, xi + 556 pp., illus. U.S. $5. 

tSummary of knowledge acquired in northern environ- 
ments from 1970 to 2000. 2001. By G. Hayem. Hydro- 
Quebec, Montreal. 1 10 pp., illus. Free. 

*Two hundred years of ecosystem and food web 
changes in the Quoddy Region, Outer Bay of Fundy. 

2002. By H. Lotze and I. Milewski. Conservation Council 
of New Brunswick, Fredericton. 

The tropical rainforest: explore the natural world of 
the rainforest swamplands and the interior. 2001. By 
G. Cheshire. Crabtree, New York. 39 pp., illus. U.S.$18. 

Visions of the wild: a voyage by kayak around Van- 
couver Island. 2001. By M. Coffey and D. Goering. 
Harbour Publishing, Madeira Park, British Columbia. 
182 pp., illus. $36.95. 

tWeird nature: an astonishing exploration of nature's 
strangest behavior. 2002. By J. Downer. Firefly Books. 
Willowdale. 156 pp., illus. Cloth $35; paper $19.95. 

Wild solutions: how biodiversity is money in the bank. 

2001. By A. Beattie and P. Ehrlich. 240 pp. U.S. $25. 95. 

Miscellaneous 

tCatalogue of meetings 1988 to 2001 Mcllwraith Field 
Naturalists. 2002. By W. W. Judd. Phelps Publishing, 
London, Ontario. 28 pp. $8. 

^Cheltenham in Antarctica. 2002. By D. M. Wilson and 
D. B. Elder. Reardon Publishing, Cheltenham, England. 
140 pp., illus. E9.99. 

tCommon and contested ground: a human and environ- 
mental history of the northwest plains. 2002. By 
T. Binnema. University of Oklahoma Press, Norman, 
xvi + 263 pp., illus. U.S.$29.95. 

*The one culture: a conversation about science. 2001. 
Edited by J. A. Labinger and H. Collins. University of 
Chicago Press, Chicago. xii+ 329 pp. U.S.$18. 

*On her own terms: Annie Montague Alexander and 
the rise of science in the American West. 2001. By 
B. R. Stein. U. California P., Berkeley, xvii + 380 pp., 
illus. U.S.$35. 



176 



The Canadian Field-Naturalist 



Vol. 116 



Books for Young Naturalists 

About amphibians: a guide for children. 200L By C. 
Sill. Peachtree, Atlanta. 40 pp., illus. U.S.$14.95. 

Animals nobody loves. 2001. By S. Simon. North-South 
Books. New York. 48 pp., illus. U.S.$15.95. 

Bears. 2001. By D. Ferte, M. Reddy, and E. D. Stoops. 
Sterling, New York. 80 pp., illus. U.S.$17.95. 

Caimans. 2001. By S. Dollar. Raintree Steck-Vaughn, 
Austin. 32 pp., illus. U.S. $23. 

Dolphins. 2001. By J. Vogel. NorthWord Press, 
Minnetonka, Minnesota. 48 pp., illus. U.S. $7. 98. 

Global Warming: the threat of earth's changing cli- 
mate. 2001. By L. Pringle. North-South Books, New 
York. 48 pp., illus. U.S.$16.95. 

Kids care for the earth. 2002. By G. Thompson. 
National Geographic Society, Washington. 32 pp., illus. 
U.S.$41.95(packof6). 

Manatees. 2001. By K. Feeney. NorthWord Press, 
Minnetonka, Minnesota. 48 pp., illus. U.S. $7.95. 



Protecting our planet : animal watch. 2001. By R. Few. 
DK Publishing, New York. 60 pp., illus. U.S. $16.95. 

Rain forest. 2001. By E. Greenwood. DK Publishing, 

New York. 48 pp., illus. U.S.$9.95. 

Recycling and reuse: our impact on the planet. 2002. 
By R. Bowden. Raintree Steck-Vaughn, Austin, Texas. 
64pp.,illus. U.S.$18.98. 

The seashore. 2001. By A. Wilkes. Kingfisher, New 
York. 32 pp., illus. U.S.$11.95. 

Sharks. 2001. By L. Evert. NorthWord Press, Min- 
netonka, Minnesota. 48 pp., illus. U.S. $7.95. 

Song birds: the language of song. 2001. By S. A. 
Johnson. Carolrhoda, Minneapolis. 48 pp., illus. 
U.S.$23.95. 

Spiny sea stars. 2001. By C. Zuchora-Walske. Lemer, 
Minneapolis. 32 pp., illus. U.S.$21.95. 

Sure-to-win science fair projects. 2001. By J. Rhatigan. 
Lark Books, New York. 128 pp., illus. U.S.$21.95. 



100 award - winning science fair projects. 2001. By G. 
Vecchione. Sterling, New York. 208 pp., illus. U.S.$21.95. 



*Assigned for review 
t Available for review. 



News and Comment 



Sea Wind: Bulletin of Ocean Voice International 5(1/2) December 2001 



This special commemorative edition combines two 
issues between its covers and is edited by Marcia Campbell 
and Noel Alfonso. Sea Wind 15(1): 1-37, was the last issue 
Don E. McAllister was working on before his death 17 
June 2001, and contains: A special notice from the editors; 
Fire Shrimp successfully cultured in Sri Lanka; A scientific 
consensus on marine reserves; Filipino fisher-folk fighting 
to benefit from marine biological riches; Free trade and 
sovereignty to protect rights of coastal communities & fish- 
ers; Satellite data helps save coral reefs; Selective fishing 
recommendations in Canada; Mapping life in the world's 
oceans; Ocean Voice International 2001 Annual General 
Meeting; Ocean Voice International Annual Report; Sea 
News. Sea Wind 15(2): 38-64, is dedicated to Don 



McAllister and contains: Memorial cover page "In memory 
of Don" a tableau of scleractinian corals by Roelof Idema; 
Ode to Grampa; Friends write in memory of Don; 
Dedication article: model citizen of the world; Curatorial 
and research contributions - Museum of Nature; Dedication 
article: Is bottom trawling scorched-earth fishing; Kid's 
korner; Tatay Don McAllister: Haribon Foundation's 
empowering partner; Ocean Voice International member- 
ship form. 

For membership rates contact Ocean Voice Inter- 
national, Box 37026, 3332 McCarthy Road, Ottawa, 
Ontario KIV OWO; Telephone (613) 721-4541; 
Fax (613) 721-4562; Office Manager at e-mail: ocean- 
voice. superaje.com; home page: http://www.ovi.ca. 



Biodiversity: Journal of Life on Earth 3(1) February 2002 



Contents: Teaming up to conserve the biodiversity of 
Western Kenys (Eusebius J. Mukhwana) — Some reflec- 
tions on the work of Don E. McAllister (Ted Mosquin) — 
Marine biodiversity of of Canada: Threats and conservation 
solutions — Black widow spiders: An outline of diversity 
(Charles D. Dondale) — The Giant Jamaican Swallowtail 
(Papilio homerus F.) — Going, going... (Peter W. Hall) — 
Forum: Omission of Biodiversity measurement and human 
population issues - shocking! (Don Kerr) — Editor's Comer 



(Ted Mosquin) — Species by species [Chocolate, 
Theobroma cacao] — Biodiversity News — Book Reviews 
— Announcements. 

The new editor of Biodiversity is Dr. Ted Mosquin; 
Catherine Ripley remains as Managing Editor. Biodiversity 
is published by The Tropical Conservancy, 94 Four 
Seasons Drive, Ottawa, Ontario K2E 7S1 Canada, e-mail: 
tropical @ synapse.net; 
URL: http://www.synapse.net/~tropical 



The Boreal Dip Net: Newsletter of the Canadian Amphibian and Reptile Conservation 
Network 6(1) December 2001 



This issue highlights the CARCNET meetings held in 
Prince Edward Island in 18-22 October 2001. [List of 
abstracts from the meeting is available on the website: 
http:\www.carcnet.ca/]. Contents: Editor's note [Kerrie 
Serben]; Report from the Chair [Christine Bishop]; CAR- 
CNET Resolution to help marine trutles in Canada; 2001 
CARCNET Award Winners: The Blue Racer Award 
[Francis R. Cook]; Silver Salamander Awards [Nature 
Trust of New Brunswick for establishment of Hyla Park 
(Fredericton) and Ducks Unlimited for their Small Marsh 
Program]; DigitalFrog International/CARCN Scholarship 



[Shana Truant: A comparison of four different salamander 
sampling techniques and a study of salamander popula- 
tions that reside in riparian reserves in Ontario's mixed- 
wood forests]; Student Platform presentations; 
Abstracts/Lecture notes from Keynote Speakers: Sherman 
Bleakney, Francis R. Cook, Michael C. James; Field trip - 
Revisiting Old Study Sites in PEI [Shana Truant]; The rare 
and secretive Sharp-tailed Snake [David Cunnington]; 
Fracine the Poster Frog ["Its not easy being green": 
Kejimkujik National Park: Peter Hope]; CARCNET List 
of Directors for 2001-2002. 



Recovery: An Endangered Species Newsletter (19) January 2002 



Published by the Canadian Wildlife Servis this issue 
contains: Recovery Highlights: Tracking turtles; Protecting 
habitat; Taking flight; Saving Ontario's mussels [David 
Wylynko]; Special Report: Endangered Species Recovery 
Fund: Supporting a diverse array of projects; Bowhead 
whale strategy releasd: Long-term conservation, ecosystem 
approach planned for eastern arctic population; Wood 
Bison recovery plan approved; Scientists launch beluga 
recovery; Caribou committee proposed; Recovering 
Newfoundland plants; Updates: Workshop held; Report 



released; Assessing soecies; Monitoring migrations: Ham 
operators assist scientists in search for wild species in 
migration; Researchers map biodiversity; New findings in 
swan research; New publications; Awards; Site seeing; 
Featured Species: Rador reveals bird's secretive behaviour 
[Marbled Murrelet]. 

Contact: Recovery, Canadian Wildlife Service, 
Environment Canada. Ottawa, Ontrio Kl A 0H3. 
Web site: www.cws-scf.ec.gc/recovery/archive.html 



77 



178 



The Canadian Field-Naturalist 



Vol. 116 



RENEW: National Recovery Plan Number 21: The Wood Bison, Bison bison athabascae 



The Recovery of Nationally Endangered Wildlife 
[Canada] has issued its "National Recovery Plan for the 
Wood Bison, Bison bison athabascae as Plan Number 21, 
October 2001. 50 pages. It was prepared by a team consist- 
ing of C. Cormack Gates, University of Calgary; Robert O. 
Stephenson, Alaska Department of Fish and Game; Hal W. 
Reynolds, Canadian Wildlife Service, Environment 
Canada; C. G. Van Zyll de Jong [deceased May 1997, for- 
merly mammalogist, Canadian Museum of Nature]; Helen 
Schwantje, British Columbia Ministry of Environment, 



Lands and Parks; Manfred Hoefs, Yukon Department of 
Renewable Resources; John Nishi, Northwest Territories, 
Wildlife and Economic Development; Normand Cool and 
Jane Chisholm, Parks Canada Agency; Adam James, 
Alberta Fish and Wildlife Service, Sustainable Resource 
Development; Bill Koonz, Manitoba Department of Natural 
Resources. Copies are available from Recovery Secretariat, 
c/o Canadian Wildlife Service, Environment Canada, 
Ottawa, Ontario KIA 0H3; Tel 819-953-1410; Fax 819- 
994-3684. 



Alberta Wildlife Status Reports numbers 37, 

The Fisheries and Wildlife Management Division of the 
Alberta Natural Resource Status and Assessment Branch, 
Alberta Environmental Protection, has released new 
Wildlife Status Reports. The Series Editor is Sherry Frazer 
(37), Sherry Frazer and Robin Outsell (38 and 39); the 
Senior Editor for (37) is Isabelle M. G. Michaud; and the 
illustrations are by Brian Huffman for all three. For a list- 
ing earlier numbers in the series, see The Canadian Field- 
Naturalist 112(1): 169 for 1-11; 113(2): 311 for 12-17; 
113(4): 686 for 18-21; 114(1): 151 for 22-25; 115(2): 390 
for 26-31; 115(3): 000 for 32-36. Recent reports issued in 
2001 are: 

37. Status of the Grizly Bear (Ursus arctos) in Alberta, by 
John L. Kansas. 43 pages. 

38. Status of the Wood Bison (Bison bison athabascae) in 



38,39 

Alberta, by Jonathan A. Mitchell and C. Cormick Gates. B I 

32 pages. 
39. Status of the Bull Trout (Salvelinus confluentus) in 

Alberta, by John R. Post and Fiona D. Johnson. 40fl| 

pages. 

For copies contact the Information Centre - Publications 
Alberta Environment/Alberta Sustainable Resource 
Development, Fish and Wildlife Division, Main Floor, 
Great West Life Building, 9920 - 108 Street, Edmonton, 
Alberta T5K 2M4 Canada (telephone: (780) 422-2079), OR 
Information Service Alberta Environment/Sustainable 
Resource Development, #100, 3115 - 12 Street NE, 
Calgary, Alberta T2E 7J2, Canada (telephone: (403) 297- 
3362); web site: http;//www3.gov.ab.ca/srd/fw/status/ 
index.html. 



Ontario Natural Heritage Information Centre Newsletter 6(1) Winter 2002 



Contents of this 20 page issue: Great Lakes Ecoregional 
Planning Project — NHIC Prepares Ontario's first "General 
Status of Wild Species" report — NHIC Assists with the 
Georgian Bay Coast Project — Kawartha Highlands 
Update — Zoology: Species at Risk Biologists Assist with 
Surveys of Threatened Fish Species — NHIC Hosts the 
First Great Lakes Odonata Meeting — Botany: 2001 
Botanical Highlights — Community Ecology: Rare 
Vegetation of Ontario: Database Cliffs of Northwestern 
Ontario — News and Notes: NHIC Establishes New 
Partner Agreement — NHIC Prepares Report on the 
Occurrence of Species at Risk in OMNR Districts — Trent 



University-NHIC Internships in Conservation Biology — 
NHiC Participates in First Pelee Island Endangered Species 
Festival — Updates to the OMNR and COSEWIC Lists of 
Species at Risk — "Bioconservation and Systematics" 
Proceedings Now Available — Farewell to Jarmo Jalava — 
Book Reviews — Focus on Gordon Wichert — Greetings 
from NIHCs New Coordinator — NHIC Staff List. 

Mailing address for Natural Heritage Information 
Centre: 300 Water Street, 2nd Floor, North Tower, P.O. 
Box 7000, Peterborough, Ontario K9J 8M5, Canada. 
Web page: http://www.mnr.gov.on.ca/MNR/nhic.html 



Marine Turtle Newsletter (95) 

The January 2002 issue, 36 pages, contains: Guest 
Editorial: The swampland of sea turtle conservation: in 
search of a philosophy — Articles: Status of the sea turtle 
trade in Alexandria's fish market — Direct carapacial 
attachment of satellite using orthopedic bioabsorbable 
mini-anchor screwws on Leatherback Turtles in Culebra, 
Puerto Rico — Notes: The reproductive status of marine 
turtles nesting in the Cayman Islands: work in progress — 
occurrence terminology for marine turtles — sea turtle 
research and conservation project in Cipara, Paria 



Peninsula, Sucre State, Venezuela: Preliminary results of 
the 2000 nesting season — Project update: University 
Progject for the study of conservation of Cuban sea turtles: 
completion of year 3 — Announcements — Book 
Reviews — News & Legal Briefs — Recent 
Publications. 

The Marine Turtle Newsletter is edited by Brendan J. 
Godley and Annette C. Broderick, Marine Turtle Research 
Group, School of Biological Sciences, University of Wales, 
Swansea, SA2 8PP Wales, United Kingdom; e-mail 



2002 



News and Comment 



179 



MTN@swan.ac.uk; Fax +44 1792 295447. Subscriptions to 
the MTN and donations towards the production of MTN 
and its Spanish edition NTM [Noticiero de Tortugas 
Marinasl should be sent to Marine Turtle Newsletter c/o 



Chelonian Research Foundation, 168 Goodrich Street, 
Lunenburg, Massachusetts 01462 USA; e-mail 
RhodinCRF@aol.com; fax + 1 978 582 6279. MTN web- 
site is: <http://www.seaturtle.org/mtn/> 



Froglog: Newsletter of the Declining Amphibian Populations Task Force (49) 



Number 49, February 2002, contains: Crisis less severe 
for Po Valley Spadefoot Pelobates fuscus insubricus 
(Vincenzo Ferri Italian DAPTF Working Group & Toads 
Project) — Effects of Habitat Disturbance on a Frog 
Community in a Mexican Dry Forest (Ireri Suazo-Ortuno) 
— Amphibian Monitoring in Africa (Cote d'lvoire, Kenya) 
and Asia (Borneo) (from Global Amphbian Diversity 
Anaysis Group) — Amphibian decline in the Kariba 
Wilderness Area, Zimbabwe (Peter Taylor) — Froglog 
Shorts — Publications of Interest. 

Froglog is the bi-monthly newsletter of the Declining 



Amphibian Populations Task Force of The World 
Conservation Union (IUCN)/Species Survival Conmiission 
(SSC) and is supported by The Open University, The 
World Congress of Herpetology, The Smithsonian 
Institution, and Harvard University. The newsletter is 
Edited by John W. Wilkinson, Department of Biological 
Sciences, The Open University, Walton Hall, Milton 
Keynes, MK7 6AA, United Kingdom; e-mail: 
daptf@open.ac.uk. Funding for Froglog is underwritten by 
the Detroit Zoological Institute, P. O. Box 39, Michigan 
48068-0039, USA. 



The 124th Annual Business Meeting of The Ottawa Field-Naturalists' Club: 7 January 2003 



The 124th Annual Business Meeting of The Ottawa 
Field-Naturalists' Club will be held in the auditorium of 
Victoria Memorial Museum (Canadian Museum of Nature), 
McLeod and Metcalfe streets, Ottawa, on Tuesday 7 
January 2003 at 7:30 p.m. (19:30 h). The Council for 2003 



will be elected at this meeting and a brief review of the 
activities during 2002 will be given, as well as a statement 
of the Club's finances. 

Ken Allison 
Recording Secretary 



Call for Nominations: The Ottawa Field-Naturalist's Club 2003 Council 



Candidates for Council may be nominated by any mem- 
ber of The Ottawa Field-Naturalist's Club. Nominations 
require the signature of the nominator and a statement of 
willingness to serve in the position for which nominated by 



the nominee. Some relevant background information on the 
nominee should also be provided. 

Fenja Brodo 
Chair, Nominating Committee 



Call for Nominations: The Ottawa Field-Naturalists' Club 2002 Awards 



Nominations are requested from members of The 
Ottawa Field-Naturalists' Club for the following: Honorary 
Membership, Member of the Year, George McGee Service 
Award, Conservation Award: Member, Conservation 
Award: Non-member, and the Anne Hanes Natural History 
Award. Descriptions of these awards have appeared in The 



Canadian Field-Naturalist 113(4): 689. With the exception 
of nominations for Honorary Member and Conservation 
Award: Non-member, all nominees must be members of 
The Ottawa Field-Naturalist Club in good standing. 

Irvin Brodo 
Chair, Awards Committee 



180 



The Canadian Field-Naturalist 



Vol. 116 



Letters to the Editor: 



Blastomycosis in Free Ranging Wolves, Canis lupus, on the North Shore of Lake Superior, 
Ontario — A response to P. Krizan 



The July-September 2000 issue of the Canadian 
Field-Naturahst 114(3): 491-493 contained an article 
titled "Blastomycosis in a Free Ranging Wolf, Canis 
lupus, on the North Shore of Lake Superior, Ontario", 
authored by Mr. Peter Krizan. We regret that Mr. 
Krizan published the article without institutional or 
contractual approval of the researchers who collabo- 
rated in the collection, analyses, and interpretation of 
the data. We view the independent publication and 
appropriation of these results as an unprofessional 
assumption of the authorship and very serious breach 
of professional ethics. For the record, the research was 
conducted under federal and provincial permits issued 
to Dr. Paul Paquet as primary investigator. By com- 
mon agreement that included Mr. Krizan, the study 
team had assigned senior authorship to Dr. Doug 
Campbell of OVC, Pathobiology, Ontario Region 
Wildlife Health Centre, University of Guelph. In 
alphabetical order, coauthors were Frank Burrows 
(Parks Canada), Anne Forshner (University of 
Alberta), Peter Krizan (Acadia University), Graham 
Neale (University of Montana), Paul Paquet 
(University of Calgary), and Keith Wade (Parks 
Canada). 

The information included in Mr. Krizan' s initial 
report is accurate but incomplete. We take this 
opportunity to expand the original note. Blasto- 
mycosis is a chronic fungal disease that affects the 
pulmonary system of humans and dogs, and occa- 
sionally other animals (Jungerman and Schwartzman 
1972; Stroud and Coles 1980; Legendre et al. 1981; 
Thiel et al. 1995). The fungus is enzootic in Min- 
nesota (Schlosser 1980) and Wisconsin (Sarosi et al. 
1979; McDonough and Kuzman 1980) but until now, 
had not been reported in other North American wolf 
populations. From 1995-1999, three adult radiocol- 
lared wolves from our study sample (n = 26) were 
diagnosed with Blastomycosis at the time of death, 
though only one (Krizan 2000) was determined to 
have died of the disease. One wolf (Spirit, male) was 
shot and another (Moon, female) killed by other 
wolves (Forshner 2000). The latter was in extremely 
poor condition and evidence at the site of her death 
suggested pack mates killed her. All three wolves 
occupied home ranges north of Pukaskwa National 
Park and were members of different packs. Recently, 
these wolves were shown to be genetically distinct 
from the Gray Wolves (C. lupus) occurring within 
Pukaskwa National Park (Wilson et al. 2000). Both 



types of wolves are known to occur in Minnesota 
and Wisconsin but the original reports of blastomy- 
cosis infections in these regions preceded genetic 
typing. Consequently, blastomycosis has not been 
unambiguously confirmed in both types of wolves. 

Literature Cited 

Forshner, S. A. 2000. Population dynamics and limitation 
of wolves {Canis lupus) in the Greater Pukaskwa 
Ecosystem, Ontario. M.Sc thesis. University of Alberta, 
Edmonton, Alberta, 129 pages. 

Jungerman, P. F., and R. M. Schwartzman. 1972. Veter- 
inary medical mycology. Lea and Febiger, Philadelphia, 
Pennsylvania, USA. 200 pages. 

Legendre, A. M., B. A. Selcer, D. F. Edwards, and R. 
Stevens. 1984. Treatment of canine blastomycosis with 
amphotericin B and ketoconazole. Journal of the Ameri- 
can Veterinary Medical Association 184: 1249-1254. 

McDonough, E. S., and J. F. Kuzma. 1980. Epide- 
miological studies on blastomycosis in the state of 
Wisconsin. Sabouraudia 18: 1373-183. 

Sarosi, G. A., M. R. Eckman, S. F. Davies, and W. K. 
Laskey. 1979. Canine blastomycosis as a harbinger of 
human disease. Annals of Internal Medicine 91: 
773-735. 

Stroud, R. K., and E. M. Coles. 1980. Blastomycosis in an 
African lion. Journal of the American Veterinary 
Medical Association 177: 842-844. 

Thiel, R. P., L. D. Mech, G. R. Ruth, J. R. Archer, and L. 
Kaufman. 1987. Blastomycosis in wild wolves. Journal 
of Wildlife Disease 23: 321-323. 

Wilson, P. J., S. Grewal, I. D. Lawford, J. Heal, A. G. 
Granacki, D. Pennock, J. B. Theberge, M. T. 
Theberge, D. Voigt, W. Waddell, R. E. Chambers, 
P. C. Paquet, G. Goulet, D. Cluff, and B. N. White. 
2000. DNA profiles of the eastern Canadian wolf and 
the red wolf provide evidence for a common evolution- 
ary history independent of the gray wolf. Canadian 
Journal of Zoology 78: 2156-2166. 

Paul C. Paquet (Conservation Science Inc. 

and University of Calgary)' 

Frank Burrows (Parks Canada 

and Lakehead University) 

Anne Forshner (University of Alberta) 

Graham Neale (University of Montana) 

Keith Wade (Parks Canada) 

Received 12 May 2001 

'Box 150, Meacham, Saskatchewan SOK 2V0 Canada; 
email: ppaquet@sk.sympatico,ca 



2002 



News and Comment 



181 



Blastomycosis in a Free ranging Wolf, Canis lupus ^ on the North Shore of Lake Superior, 
Ontario — A reply to P. Paquet et al. 



Paquet et al. report interesting results that add 
valuable information to the original note by Krizan 
2000 and suggest that Blastomycosis is more preva- 
lent than was originally thought and may not always 
be fatal. My note (Krizan 2000) was one finding of 
my Masters thesis that was started in 1993 and was 
completed and defended in 1997 (Krizan 1997). It 
should be noted that the additional data provided by 
Paquet et al. were collected after the completion of 
my thesis. 

In both publications (Krizan 2000, 1997), I 
acknowledged the members and agency involved in 
the co-operative project for their role and participa- 
tion. Parks Canada (Pukaskwa National Park) sup- 
ported the Pukaskwa National Park Predator Prey 
Process Project (P5) through logistical support. In 
addition, many volunteers and other researchers 
working on different aspects of this project were 
involved to different degrees; those who had con- 
tributed directly to the collection of data were men- 
tioned by name in the acknowledgment sections of 
Krizan 1997 and 2000. 1 once again acknowledge the 
role of some of the co-authors of Paquet et al. in the 
overall project and their recent contribution. 

I was one of three students completing a Masters 
thesis from 1993 to 1997. The only agreement that I 
had with Parks Canada and/or P. Paquet, who was 
the principal investigator contracted by Parks 
Canada was that Parks Canada would supply the 
logistical support for my project and in return, I 
would collect, analyze and publish my results and 
make my thesis and raw data available to the 
Pukaskwa National Park office. I have fulfilled all of 
those responsibilities. During 1993 to 1997 I was an 
independent student with no contractual agreements 
with any of the co-authors of Paquet et al. I was the 
only one that analyzed data that was used in the 
Krizan 2000 publication except for the necropsy that 
was done by Dr. Doug Campbell (as was acknowl- 
edged). I had discussed Dr. Campbell's necropsy 
results with him and he provided me with relevant 
literature. To the best of my knowledge, the research 
results of the Masters thesis are owned by copyright 



and are not subject to institutional or contractual 
consent. My research interest was to describe the 
travel patterns of Wolves and how these movements 
were affected by human presence, activity, and land- 
scape differences in and outside of Pukaskwa 
National Park. The case of blastomycosis was inter- 
esting in that infection by this fungus was probably 
due to the fact that this Wolf was a lone animal that 
exposed itself to areas by travelling in environments 
that may have had a higher probability of exposure 
to the fungal spores. Also interesting was the fact 
that the travel distances between locations dimin- 
ished as the disease affected the ability of the Wolf 
to travel. To the best of my knowledge, I was the 
only person on the P5 project team that was interest- 
ed in the spatial use of Wolves. As with any project, 
there are many people that are involved to various 
degrees and I've acknowledged those who had con- 
tributed directly to the collection of data. 

Two of the co-authors of Paquet et al. were also 
students working on their theses of various but dif- 
ferent topics. I am confused about the role and co- 
authorship of Miss Forshner. I met Miss Forshner 
once as I was leaving the project. I had never worked 
with her nor did I ever discuss authorship of publica- 
tions. The implied consent to co-author any publica- 
tion in the order that Paquet et al. suggest is incor- 
rect. 

Literature Cited 

Krizan, P. 1997. The effects of human land development, 
landscape characteristics, and prey density on the spatial 
distribution of Wolves {Canis lupus) on the north shore 
of Lake Superior, Ontario. M.Sc. thesis, Acadia Univer- 
sity, Wolfville, Nova Scotia. 108 pages. 

Krizan, P. 2000. Blastomycosis in a free ranging lone 
Wolf, Canis lupus, on the north shore of Lake Superior, 
Ontario. Canadian Field-Naturalist 1 14: 491-493. 

Peter Krizan 
P.O. Box 1 167 Iqaluit, Nunavut XOA OHO Canada 

Received 5 December 200 1 



Advice for Contributors to The Canadian Field-Naturalist 



Content 

The Canadian Field-Naturalist is a medium for the publi- 
cation of scientific papers by amateur and professional natu- 
ralists or field-biologists reporting observations and results 
of investigations in any field of natural history provided that 
they are original, significant, and relevant to Canada. All 
readers and other potential contributors are invited to submit 
for consideration their manuscripts meeting these criteria. 
The journal also publishes natural history news and com- 
ment items if judged by the Editor to be of interest to read- 
ers and subscribers, and book reviews. Please correspond 
with the Book Review Editor concerning suitability of 
manuscripts for this section. For further information consult: 
A Publication Policy for the Ottawa Field-Naturalists' Club, 
1983. The Canadian Field-Naturalist 97(2): 231-234. 
Potential contributors who are neither members of The 
Ottawa Field-Naturalists' Club nor subscribers to The 
Canadian Field-Naturalist are encouraged to support the 
journal by becoming either members or subscribers. 

Manuscripts 

Please submit, to the Editor, in either English or French, 
three complete manuscripts written in the journal style. 

The research reported should be original. It is recommended 
that authors ask qualified persons to appraise the paper 
before it is submitted. All authors should have read and 
approved it. Institutional or contract approval for the publi- 
cation of the data must have been obtained by the authors. 
Also authors are expected to have complied with all perti- 
nent legislation regarding the study, disturbance, or collec- 
tion of animals, plants or minerals. The place where voucher 
specimens have been deposited, and their catalogue num- 
bers, should be given. Latitude and longitude should be 
included for all individual localities where collections or 
observations have been made. 

Print the manuscript on standard-size paper, double- 
space throughout, leave generous margins to allow for 
copy marking, and number each page. For Articles and 
Notes provide a bibliographic strip, an abstract and a list of 
key words. Generally, words should not be abbreviated but 
use SI symbols for units of measure. The names of authors 
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manuscripts or other papers involving nomenclatural prob- 
lems. "Standard" common names (with initial letters capi- 
talized) should be used at least once for all species of high- 
er animals and plants; all should also be identified by scien- 
tific name. 

The names of journals in the Literature Cited should be 
written out in full. Unpublished reports should not be cited 
here but placed in the text or in a separate Documents Cited 
section. List the captions for figures numbered in arable 
numerals and typed together on a separate page. Present the 
tables each titled, numbered consecutively in arabic numer- 
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the text the places for the figures and tables. 

Check recent issues (particularly in literature cited) for 
journal format. Either "British" or "American" spellings are 



acceptable in English but should be consistent within one 
manuscript. The Oxford English Dictionary, Webster's 
New International Dictionary and le Grand Larousse 
Encyclopedique are the authorities for spelling. 

Illustrations 

Photographs should have a glossy finish and show sharp 
contrasts. Photographic reproduction of line drawings, no 
larger than a standard page, are preferable to large origi- 
nals. Prepare line drawings with India ink on good quality 
paper and letter (don't type) descriptive matter. Write 
author's name, title of paper, and figure number on the 
lower left comer or on the back of each illustration. 

Reviewing Policy 

Manuscripts submitted to The Canadian Field-Naturalist 
are normally sent for evaluation to an Associate Editor 
(who reviews it or asks another qualified person to do so), 
and at least one other reviewer, who is a specialist in the 
field, chosen by the Editor. Authors are encouraged to sug- 
gest names of suitable referees. Reviewers are asked to give 
a general appraisal of the manuscript followed by specific 
comments and constructive recommendations. Almost all 
manuscripts accepted for publication have undergone revi- 
sion — sometimes extensive revision and reappraisal. The 
Editor makes the final decision on whether a manuscript 
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tain the scientific quality, content, overall high standards 
and consistency of style, of the journal. 

Special Charges — Please take note 

Authors must share in the cost of publication by pay- 
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to $80 per page for tables (depending on size). Repro- 
duction of color photos is extremely expensive; price quo- 
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Reprint order forms are included when galley proofs are 
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Reprints 

An order form for the purchase of reprints will accom- 
pany the galley proofs sent to the authors. 

Francis R. Cook, Editor 

RR 3 North Augusta, 

Ontario KOG IRO, Canada 



182 



TABLE OF CONTENTS (concluded) 

(^olf, Canis lupus, response to domestic sled dog, Canisfamiliaris, 
activities in central Yukon Gerald W. Kuzyk and Kristin M. Kuzyk 125 

irst record of an Eastern Coyote, Canis latrans, in Labrador 

ToxY E. Chubbs and Frank R. Phillips 127 

ong distance movement by a Coyote, Canis latrans, and a Red Fox, Vulpes vulpes, 
in Ontario: Implications for disease-spread Richard C. Rosatte 129 

redation of Wolves, Canis lupus, on Wolverines, Gulo gulo,and an American Marten, 
Martes americana., in Alaska Kevin S. White, Howard N. Golden, 

Kris J. Hltvidertmark, and Gerald R. Lee 132 

leek's Halfbeak, Hyporhamphus meeki, and Flying Gurnard, Dactylopterus volitans, 
captured in the Annapolis Basin, Nova Scotia A. Jamie F. Gibson and Ransom A. Myers 134 

pparent capture myopathy in Hoary Bats, Lasiurus cinereus: A cautionary note 

Thomas S. Jl^g, Un D. Thompson, M. Brian C. Hickey, and Roger D. Titman 136 

lobbing Black-billed Magpie, Pica hudsonia, killed by Cooper's Hawk, Accipiter cooperii 

Geoffrey L. Holroyd 137 

reeding season of Wolves, Canis lupus, in relation to latitude L. David Mech 139 

[istorical Feature Article 

anada and the "buffalo". Bison bison: A tale of two herds W. A. Fuller 141 

ook Reviews 

oology: Birds, Mammals & Reptiles of the Galapagos Islands — Wisconsin Fishes 2000: Status and 
Distribution — Marine Mammals of the Pacific Northwest: A Concise and Comprehensive 
Waterproof Guide — Manitoba Birds — Heron Conservation — The Field Guide to Birds of 
Australia — Katydids and Bush-Crickets: Reproductive Behavior and Evolution of the 
Tettigoniideae — A Manual for Wildlife Radio Tagging — Bird Census Techniques — Flying 
Foxes: Fruit and Blossom Bats of Australia 1 60 

otany: Guide to Standard Floras of the World: An Annotated Geographically Arranged Systematic 
Bibliography of the Principal Floras, Enumerations, Checklists, and Chorological Atlases of 
Different Areas, Second Edition 168 

nvironment: Their Father's Work: Casting Nets with the World's Fishermen — AAAS Atlas of 
Population and Environment — Making Better Environmental Decisions — Scientific Method for 
Ecological Research 1 69 

liscellaneous: The Last Great Sea: A Voyage through Human and Natural History of the 
North Pacific Ocean 172 

ew Titles 174 

lews and Comments: 

?a Wind: Bulletin of Ocean Voice International 5(1/2) December 2001 — Biodiversity: Journal of Life 
on Earth 3(1) February 2002 — The Boreal Dip Net: Newsletter of the Canadian Amphibian and 
Reptile Conservation Network 6(1) December 2001 — Recovery: An Endangered Species 
Newsletter (19) January 2002 — RENEW: National Recovery Plan Number 21: The Wood Bison. 
Bison bison athabascae — Alberta Wildlife Status Reports numbers 37, 38, 39 — Ontario Natural 
Heritage Information Centre Newsletter 6(1) Winter 2002 — Marine Turtle Newsletter (95) — 
Frogleg: Newsletter of the Dectining Amphibian Population Task Forces (49) — Notice of the 124lh 
Annual Business Meeting of The Ottawa Field-Naturalists' Club: 7 January 2003 — Call for 
Nominations: The Ottawa Field-Naturalist's Club 2003 Council — Call for Nominations: The 
Ottawa Field-Naturalists' Club 2002 Awards 1 77 

etters to the Editor: Blastomycosis in free-ranging Wolves, Canis lupus, on the north shore of Lake 
Superior. Ontario: A response to P. Krizan; A reply to Paquet et al. 1 80 

idvice to Contributors 1 82 

(ailing date of the previous issue 1 15(4): 9 August 2002 



THE CANADIAN FIELD-NATURALIST 



Volume 116, Number 1 



Articles 

Spread and disappearance of the Greater Prairie-Chicken, Tympanuchus cupido, 
on the Canadian prairies and adjacent areas C. Stuart Houston 

Status of Common Eiders, Somateria mollissima, nesting in the 
Digges Sound region, Nunavut J. Mark Hipfner, H. Grant Gilchrist, 

Anthony J. Gaston, and David K. Cairns 

Flathead Chubs, Platygobio gracilis, in the upper Missouri River: The biology of a species 
at risk in an endangered habitat ---"m^M. ^-S^annon J. Fisher, David W. Willis, 

o^ .i^^^^^s^^iQH^^ M. Olson, and Steven C. Krentz 

Abundance and distribution of breeding waterfowl in the Great Clay Belt of northern Ontario 

R. Kenyon Ross, Kenneth F. Abraham, Ted R. Gadawski, 
Robert S. Rempel, T. Shane Gabor, and Ron Maher 

Resilience of Foothills Rough Fescue, Festuca campestris, rangeland to wildfire 

Edward W. Bork, Barry W. Adams, and Walter D. Willms 

Survival, fates, and success of transplanted Beavers, Castor canadensis, in Wyoming 

Mark C. McKinstry and Stanley H. Anderson 

Songbird community composition versus forest rotation age in Saskatchewan boreal 
mixedwood forest Enid E. Gumming and Antony W. Diamond 

Status of Redside Dace, Clinostomus elongatus, in the Lynde Creek and Pringle Creek 
watersheds of Lake Ontario Jeff J. Anderson 

Aquatic leaves and regeneration of last year's staw in the arctic grass, Arctophila fulva 

Susan G. Aiken and Rosemary A. Buck 

Records of Northern Mockingbird, Mimus polyglottos, occurrences in North Dakota 
during the twentieth century Lawrence D. Igl and Ron E. Martin 

Diets of Northern Flying Squirrels, Glaucomys sabrinus, in southeast Alaska 

Sanjay Pyare, Winston P. Smith, Jeffrey V. Nicholls, and Joseph A. Cook 

Nesting activities of an Eastern Spiny Softshell Turtle, Apalone spinifera 

Claude Daigle, Patrick Galois, and Yves Chagnon 

Long-distance movements by female White-footed Mice, 
Peromyscus lecopus, in extensive mixed-wood forest Thomas J. Maier 

Recent trends in stem numbers in Goldenseal, Hydrastis canadensis, populations at the 
northern limit of its range Adrianne Sinclair and Paul M. Catling 

A comparison of techniques for assessing amphibian assemblages on streams in the 

Cynthia A. Paszowski, Garry Scrimgeour, 
Beverly A. Gingras, and Sharon Kendall 



I 



western boreal forest 



Notes 

A rare leucistic Spiny Dogfish, Squalus acanthias, from the Bay of Fundy, Nova Scotia 

Brian W. Coad and John Gilhen 

Anomalies in incisor wear of American Elk, Cervus elaphus, in the French River delta, 
Ontario J. Hamr, F. F. Mallory, I. A. Filion, G. S. Brown, and M. A. Jost 

First record of the Hoary Bat, Lasiurus cinereus (Chiroptera: Vespertilionidae), 
from Prince Edward Island Donald F. McAlpine, Frances Muldoon, 

and Alexander I. Wandeler 



SMrTHSONIAN INSTrtUTION LIBRA 



ISSN 0008-3550 




3 9088 01226 61 




'j!^ The CANADIAN 
FIELD-NATURALIST 



<j 



Published by THE OTTAWA FIELD-NATURALISTS' CLUB, Ottawa, Canada 




W" 



/olume 116, Number 2 



April-June 2002 



The Ottawa Field-Naturalists' Club 

FOUNDED IN 1879 

Patrons 

Her Excellency The Right Honourable Adrienne Clarkson, C.C., C.M.M., CD. 

Governor General of Canada 
His Excellency John Ralston Saul, C.C. 

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Cover: Logan Glacier and the St. Elias Mountains in Wrangell-St.Elias National Park and Preserve, Alaska. The site in tha | 
foreground is the locality of A rabis calderi G A. Mulligan, which is new to Alaska, and also the collection locality or 
the mustards Braya glabella Richardson ssp. glabella, Draba cinera Adams, D. incerta Payson, D. macounii O. E. 
Schultz, D. oligosperma Hook, and Lesquerella arctica (Wormsk.) S. Wats. ssp. arctica, all of which were range! i 
extensions at this site. Silene menziesii Hook., Carex filifolia Nutt., and Danthonia intermedia Vasey were other* ' 
range extensions from this locality. See article on notable vascular plants form Alaska by Mary B. Cook and Carl A. 
Roland, pages 192-304. 



The Canadian Field-Naturalist 



Volume 116, Number 2 



April-June 2002 




n Elk, 



Winter Habitat Selection at Three Spatial S^ 
Cervus elaphus, in West-Central Alberta 

Paul F. Jones, i and Robert J. Hudson -v - * 

i^BRARlES 

Department of Renewable Resources, University of Alberta, Edmonton, Alberta T6Lr Ihi, L'ana5a 
iPresent address: Alberta Conservation Association, 2nd Floor, YPM Place, 530 - 8th Street S., Lethbridge, Alberta 
TIJ 2J8 Canada; E-mail: paul.jones@gov.ab.ca 

Jones, Paul P., and Robert J. Hudson. 2002. Winter habitat selection at three spatial scales by American Elk, Ceniis ela- 
phus, in west-central Alberta. Canadian Field-Naturalist 116(2): 183-191. 

Selection by American Elk (Cervus elaphus) at the landscape (2"^^ order), stand (3'^'^ order), and site level (4"^ order) was 
examined in west-central Alberta from 1 December 1994 to 21 March 1995. At the landscape level, elk home ranges had a 
lower mean road density, but greater mean unimproved access density. Elk home ranges had smaller mean patch sizes and 
greater patch density. There was no significant selection on the basis of food and cover composition. At a stand level, elk 
used the grass/meadow habitat more than expected while all other habitat types were used in proportion to their availabili- 
ty. At a site scale, we attempted to distinguish habitat use for feeding and bedding. Feeding sites had a lower mean percent 
canopy closure, percent shrub cover, stem density and tree height, but a higher mean percent grass cover than bedding 
sites. Feeding sites were significantly closer to unimproved access than were bedding sites. Feeding sites had more grass 
cover and significantly less canopy closure, percent shrub cover, spruce, pine, and fir in the canopy, and lower tree height 
and stem density than the control plots. They were located significantly closer to unimproved access and farther from hid- 
ing cover. Bedding sites had a higher percent grass cover, less spruce, pine and fir in the canopy, and a lower stem density 
than the control plots. In terms of spatial attributes, elk exhibited no selection when choosing bedding sites. The need to 
examine habitat selection patterns at more than one scale is discussed 

Key words: Cervus elaphus, American Elk, habitat, landscape, site, spatial scale, stand. Alberta. 



Resource selection is an adaptive response of ani- 
mals attempting to meet their life requirements and 
ultimately to increase fitness (Johnson 1980; Thomas 
and Taylor 1990; Manly et al. 1993). Choice of habi- 
tat is governed by a variety of factors including ener- 
gy requirements, thermal conditions, reproductive 
requirements, and intra- and interspecific competi- 
tion (Litvaitis et al. 1986; Morrison et al. 1992). 

Johnson (1980) defined four levels or scales of 
habitat selection: (1) selection of a physical or geo- 
graphical range, (2) selection of a home range within 
that geographical range, called landscape selection, 
(3) selection of habitat components within the home 
range, termed stand selection and (4) selection of 
food items at a particular feeding site. Lofroth 
(1993) expanded Johnson's (1980) fourth-order 
selection to include habitat components required for 
such activities as mating, calving, escaping predators 
and for shelter. We define this level of selection as 
site selection. 

There are several possible outcomes to habitat 
studies conducted at multiple scales. Selection main- 
ly at the landscape level is important in several 
species such as the Spotted Owl {Strix occidentalis). 



Hansen et al. (1993) concluded that landscape level 
habitat features describing old growth forests were 
the best predictors of spotted owl abundance. At the 
other end of the spectrum, a species may select habi- 
tat characteristics mainly at a site level. This may be 
the case for generalists who are able to utilize differ- 
ent local conditions found in a variety of landscapes 
(Pearson 1993). The probable outcome of multiple 
level habitat selection is a species selecting habitat 
characteristics at more than one scale. Lofroth 
(1993) determined that Marten {Martes americcma) 
selected mature to old growth serai stage habitats 
within their home range (landscape level), while 
showing no preference for any habitat type, but 
avoided young serai stages, xeric habitat types and 
wetlands (stand level). At the site level. Marten 
selected structural characteristics that were different 
from the prevailing characteristics of the habitat type 
(Lofroth 1993). Selection at more than one scale 
may be complementary to each other or selection at 
one level may explain why it occurred at a higher 
level (Pedlar etal. 1997). 

Previous research has suggested that habitat selec- 
tion by elk {Cervus cUiphus) has been governed by 



83 



184 



The Canadian Field-Naturalist 



Vol. 116 



forage, thermal cover and hiding cover requirements 
(Hersey and Leege 1976; Lyon 1979; Gates and 
Hudson 1981; Irwin and Peek 1983; Marcum and 
Scott 1985; Edge et al. 1987; McCorquodale 1987). 
These studies have examined selection at one specif- 
ic level. We examined resource use by elk at the 
landscape (2"*^ order), stand (3'^'^ order) and site level 
(4'^ order). The central hypothesis was that elk select 
habitats for foraging, thermal cover and hiding cover 
at more than one scale. Our first objective was to 
determine if elk selected home ranges based on habi- 
tat components. Our second objective was to deter- 
mine if elk selected specific habitat types within 
their home ranges. Our third objective was to deter- 
mine whether elk selected habitat characteristics to 
meet different life requisites (i.e., feeding and bed- 
ding), and whether or not elk select specific sites 
within the immediately available area on the basis of 
non-spatial and spatial attributes. 

Study Area 

The study was conducted in west-central Alberta 
along the north and south facing slopes of the 
Athabasca River Valley near Hinton, Alberta 
(53°25 N, 117°35'W). The area falls within the 
Lower Foothills Natural Subregion and rises in ele- 
vation from 500m to 1150m (Beckingham et al. 
1996). The winter climate of the Lower Foothills 
Subregion is prone to warming by chinook winds 
(Beckingham et al. 1996). Mean monthly tempera- 
ture in winter (November through February) is 
-7.8°C with a total annual precipitation of 60 mm 
(Beckingham et al. 1996). During the study (Decem- 
ber 1994 to March 1995) the m.ean monthly tempera- 
ture was -7.6°C, with the study area receiving 
35 mm of precipitation, predominantly as snow. 

Vegetation in the study area was primarily mixed- 
wood forest, some of which were stands regenerating 
after clearcut logging. The area is dominated by 
mature mixedwood stands of Trembling Aspen 
(Populus tremuloides), Lodgepole Pine (Pinus contor- 
ta). White and Black Spruce (Picea glauca and P. 
mariana respectively), and Balsam Poplar {Populus 
balsamifera). Low-bush Cranberry {Viburnum edule). 
Prickly Rose {Rosa acicularis), Green Alder {Alnus 
crispa), Canada Buffalo-berry {Shepherdia canaden- 
sis). Marsh Reed Grass {Calamagrostis canadensis), 
and Hairy Wild Rye {Elymus innovatus) are common 
understorey species (Beckingham et al. 1996). A few 
large meadows also were present in the study area. 

Methods 

Study Animals and Site Location 

From June 1 993 to September 1 994 elk were cap- 
tured using up to eight collapsible clover traps 
(Thompson et al. 1989) baited with salt blocks or 
alfalfa. Traps were placed in areas of known elk use 
and checked twice daily. Captured elk were fitted 
with a Lotek LMRT-4 radio collar containing a mor- 



tality sensor and a head up/head down activity sen- 
sor. All animals captured were handled with care in 
accordance to the principles and guidelines outlined 
by the Canadian Council of Animal Care. Twelve 
animals (10 cows and 2 bulls) were tracked through- 
out 24-hr days during the winter of 1994/1995. Elk 
were located at least twice a week from 1 December 
1994 to 21 March 1995, subject to field restrictions. 
Animals were located after 30 November 1994 to 
lessen the effect of hunting on habitat selection, 
while the 21 March 1995 date corresponded to a 
shift back to spring/summer range. The necessity of 
obtaining approximately 20 locations per animal 
over the winter period prohibited a complete ran- 
domized design. To reduce autocorrelation of suc- 
cessive locations (Swihart and Slade 1985), individ- 
ual animals were located at least 72 hours apart 
(mean = 95.5 hours). 

A Telonics TR-2 receiver with scanner and three 
element yagi antenna received signals from radio col- 
lared elk. Locations were taken from the ground from 
fixed bearing sites, with animal locations determined 
from 3-5 intersecting compass bearings. The interval 
between bearings was usually less than 25 minutes. 
We attempted to minimize distance between receiver 
and collared elk without disturbing the elk. We tried 
to visit each fix within 24 hours to confirm elk activi- 
ty (feeding, bedding, or travelling) at that point (84% 
confirmed). Additional feeding and bedding sites 
were determined by visual sightings of non-collared 
elk feeding or bedded, and the discovery of feeding 
or bedding sites while performing radio telemetry. 
Only one bed or feeding site was measured per group 
of animals. Universal Transverse Mercator (UTM) 
coordinates for all locations were determined using a 
Trimble Global Positioning System (GPS) unit 
(Trimble Navigation 1992a) and differentially cor- 
rected using the software Pathfinder (Trimble 
Navigation 1992b). 

Landscape Level Selection 

Only study animals that were located > 18 times 
during the winter were included for analysis at the 
landscape level (n = 9; 8 cows and 1 bull). The 95% 
adaptive kemal (ADK) home ranges were estimated 
for nine of the 12 collared elk using the program 
CALHOME (Kie et al. 1996). The 95% ADK UTM 
coordinates for the nine elk home ranges were 
imported into ARC/INFO (ESRI 1990) for spatial 
analysis. Weldwood of Canada's (Hinton Division) 
forest cover (Phase III data), roads, unimproved 
access (seismic lines, cut-lines, power lines and 
pipeline right-of-ways) and forest harvest data were 
summarized for each home range. To determine 
availability of habitat at the landscape level, nine 
random circular home ranges equal to the mean area 
of the nine elk home ranges were created in 
ARC/INFO (ESRI 1990). We termed these random 
circular home ranges available home ranges (AHR). 



2002 



Jones and Hudson: Winter Habitat Selection by American Elk 



185 



Each AHR was randomly placed but had to fall com- 
pletely within the study area boundary. The forest 
cover, roads, unimproved access, and forest harvest 
data were clipped and summarized for each AHR (n 
for available = 9). 

Stands were classified using the Phase III cate- 
gories of the Alberta Forest Inventory method 
(Alberta Forest Service 1981) into food, thermal 
cover and hiding cover separately for each home 
range and AHR. A stand with canopy closure no 
greater than 50% was classified as food. Thermal 
cover was a stand with canopy closure of 5 1 % or 
greater and a tree height of 12 m or greater. Hiding 
cover was defined as a stand with canopy closure of 
50% or greater. These definitions were based on 
standard definitions for elk habitat (Thomas et al. 
1979) with modifications to represent available local 
habitat types. A stand could be classified into more 
than one type under the assumption that elk utilize 
certain stands for more than one purpose. For exam- 
ple a stand could be used for both thermal and hiding 
cover. 

For each home range and AHR, the road density, 
unimproved access density, patch density, and mean 
patch size were calculated. Road density was calcu- 
lated as the kilometers of permanent roads found 
within an elk's home range or AHR, while unim- 
proved access density was calculated as the kilome- 
ters of seismic lines, cut lines, power lines and gas 
line rights-of-way found within an elk's home range 
or AHR. Patch density was defined as the number of 
forest cover stands (> 1 ha) per km^, while mean 
patch size was the mean size of forest cover stands. 
Finally the percent of each home range and AHR 
previously logged (between 1957 and 1985) was 
determined. To evaluate habitat selection at the land- 
scape level, we compared the composition of the elk 
home ranges to the AHR using the Kruskal-Wallis 
test. 

Stand Level Selection 

Locations contained within the 95% ADK home 
range for each elk (n = 9) were classified as 1 of 10 
habitat types: grass/meadow, regenerating cut, water, 
sapling, pole, open mature, closed mature, open old 
growth, closed old growth or other. Open stands had 
a canopy cover less than 50% while closed stands 
had a canopy greater than 49%. White and Garrott 
(1990) recommended determining habitat selection 
at an individual level because different animals may 
use the habitat differently. However, the number of 
relocations for each animal was inadequate for anal- 
ysis at an individual level (Neu et al. 1974). There- 
fore, locations from each collared animal were 
pooled. Habitat availability was determined by aver- 
aging the percent composition of each habitat type 
within each elk's home range. A chi-square test was 
used to determine if elk used each habitat type in 
proportion to its availability (Neu et al. 1974). If the 



null hypothesis that elk were using the habitat in 
proportion to its availability was rejected, a 
Bonferroni Z statistic was calculated for each habitat 
type to determine if it was used more or less than 
expected (Neu et al. 1974). 

Site Level Selection 

Within a home range, specific sites are used for 
specific purposes. For all feeding and bedding sites, 
a number of vegetative and spatial variables were 
determined. Visual estimates (percent) of grass 
cover, herb cover, and shrub cover were assessed in 
a 0.04 ha circular plot (radius 1 1.3 m) centered at the 
site location. Also within the 0.04 ha plot the num- 
ber of trees were counted and classified to determine 
percent spruce, pine and fir in the canopy and tree 
density (scaled to #/ha X 1000). Average tree height 
was determined by averaging the height of five trees, 
representing the stand, as determined using a cli- 
nometer (Bessie 1995). Canopy closure was estimat- 
ed at the center of the plot using a spherical den- 
siometer. All measurements were taken between 
June 1995 and September 1995, except percent can- 
opy closure, which was measured in May. 

Spatial variables were estimated from maps pro- 
duced by overlaying the feeding and bedding loca- 
tions with Weldwood of Canada's (Hinton Division) 
forest cover, road and seismic line data using 
ARC/INFO software (ESRI 1990). Distances to 
major roads, minor roads, unimproved access, forage 
stands, thermal cover stands and hiding cover stands 
were determined. All variables were estimated in 
meters. Major roads were defined as any hard 
packed road, open year round and maintained, while 
minor roads were defined as roads that are not main- 
tained year round and are usually only accessible by 
4X4 vehicle during the winter. Unimproved access 
was an inclusive term for seismic lines, cut lines, 
power lines and gas line right-of-ways. A 1-way 
ANOVA was used to determine if the mean spatial 
and non-spatial attributes at elk feeding sites were 
significantly different than those found at elk bed- 
ding sites. 

To determine if elk selected particular sites for 
feeding and bedding, we located a paired control plot 
300 m from the feeding or bedding site, in a random 
direction (Brown 1994). The non-spatial and spatial 
variables estimated at the feeding and bedding sites 
were also assessed at the control plots. For each non- 
spatial and spatial attribute, the means for the feed- 
ing and bedding sites were compared to the means at 
their paired control plot using a paired t-test. All 
tests were preformed using SPSS (Norusis 1993). 

Results 

Limd scape Level Selection 

The mean 95% ADK home range size for the nine 
collared elk was 23 km- (range: 12-53 km-). Four 
landscape variables were significantly different 



186 



The Canadian Field-Naturalist 



Vol. 116 



Table 1 . Mean attributes for elk home ranges (n = 9) and available home ranges ( AHR) (n = 9) for the winter 
of 1994-1995 in west-central Alberta. 





Elk Home Range"* 


AHR" 






M + SE 


Mean + SE 


P Value*^ 


Habitat Variables 








Food (%) 


49.22 ±1.93 


48.47 ± 6.95 


0.453 


Thermal Cover (%) 


43.28 ±2.17 


41.92 ±6.17 


0.427 


Hiding Cover (%) 


88.16 ±1.28 


82.21 ±4.88 


. 0.508 


Harvested (%) 


10.50 ±3.37 


7.92 ±4.10 


0.269 


Density Variables 








Road Density (km/km^) 


0.32 ± 0.05 


0.82 ±0.22 


0.005 


Unimproved Access Density (km/km^) 


3.18±0.16 


2.48 ± 0.24 


0.038 


Patch Density(#/km2) 


42.99 ±6.12 


21.67 ±4.85 


0.012 


Mean Patch Size (km^) 


0.03 ± 0.004 


0.06 ± 0.009 


0.007 



^ Mean home range size = 23 km^ 

'' Available home range size = 23 km- 

'^ P values are the results from Kruskal-Wallis tests. 



between the elk home ranges and the AHR (Table 
1). Elk home ranges had a lower mean road density, 
but a greater mean unimproved access density than 
the AHR. They also had smaller mean patch size and 
a greater patch density. There was no significant dif- 
ference in the composition of the home ranges and 
AHR in terms of food and cover composition. 

Stand Level Selection 

For all elk home ranges combined (n = 9), the 
dominant cover type was closed old growth while 
sapling and water (i.e., lake and river) were the least 
prevalent cover types (Table 2). Grass/meadow 
accounted for 7% of the combined home ranges, 
while regenerating cuts accounted for 10%. The 
grass/meadow cover type was used more than 
expected, while all other habitat types were used in 
proportion to their availability (Table 2). 

Site Level Selection 

Five vegetative variables were significantly dif- 
ferent between the 96 elk feeding and 62 bedding 



sites (Table 3). Compared to bedding sites, feeding 
sites had a lower mean percent canopy closure, per- 
cent shrub cover, tree density and tree height, but a 
higher mean percent grass cover. Feeding sites were 
significantly closer to unimproved access than bed- 
ding sites (Table 3). Although this comparison high- 
lighted use of specific resources for specific activi- 
ties, we also attempted to determine selection 
relative to availability. 

All of the non-spatial habitat attributes were sig- 
nificantly different between the feeding sites and 
control plots, except for percent herbaceous cover 
(Table 4). Feeding sites had significantly less canopy 
closure, and less spruce, pine, and fir in the canopy 
than the control plots. Feeding sites had a signifi- 
cantly lower tree height and stem density than the 
control plots. Feeding sites had a greater percent 
grass cover and less percent shrub cover than the 
control plots. Only two of the five spatial variables 
were significantly different between elk feeding sites 
and the control plots (Table 4). Feeding sites were 



Table 2. Habitat use at a stand level by elk (n = 9) in west-central Alberta during the winter of 1994-1995. 







Observed 


Expected 




Bonferroni 95% Confidence 


Habitat 


Available {%f 


(n) 


(n) 


X2 


Interval 


Use 


Grass / Meadow 


1 


43 


13 


64.94 


0.137-0.283 


More 


Regenerating Cut 


10 


21 


20 


0.03 


0.048-0.157 


Equal 


Water 


2 





4 


3.83 


0.000 - 0.000 


No Use 


Sapling 


3 


4 


5 


0.39 


-0.005 - 0.044 


Equal 


Pole 


10 


12 


20 


3.22 


0.016-0.101 


Equal 


Open Mature 


7 


18 


16 


0.37 


0.037-0.139 


Equal 


Closed Mature 


16 


40 


33 


1.56 


0.124-0.266 


Equal 


Open Old-growth 


11 


15 


24 


3.17 


0.026-0.120 


Equal 


Closed Old-growth 


26 


39 


53 


3.77 


0.120-0.126 


Equal 


Other 


8 


13 


17 


0.85 


0.020-0.107 


Equal 



"Pooling all elk locations and defining the area of availability as the mean of each habitat type found within each elk's 95% 
ADK home range 



2002 



Jones and Hudson: Winter Habitat Selection by American Elk 



187 



Table 3. Mean non-spatial and spatial attributes at elk feeding (n=96) and bedding sites (n=62) in west-central 
Alberta durins the winter of 1994-1995. 







Elk Feeding Sites 


Elk Bedding Sites 








M±SE 


Mean ± SE 


P Value'-" 


Non-spatial Variables 










Canopy Closure (%) 




3 1.34 ±3.45 


64.68 ± 4.03 


<0.001 


Grass Cover (%) 




59.73 ±3.37 


34.95 ± 3.30 


<0.001 


Herb Cover (%) 




19.00 ± 1.16 


16.85 ±0.89 


0.185 


Shrub Cover (%) 




15.15 ± 1.49 


27.37 ±1.76 


<0.001 


Spruce, Pine, & Fir in the canopy (%) 


28.17 ±4.20 


38.53 ±5.07 


0.120 


Stem Density (# / ha X 


1000) 


1.31 ± 0.19 


2.03 ±0.17 


0.008 


Tree Height (m) 




5.57 ±0.72 


13.38 ±0.86 


<0.001 


Spatial Variables 










Major Roads 




1093 ± 66.06 


1244 ±87.46 


0.164 


Minor Roads 




601 ± 70.05 


763 ± 69.90 


0.121 


Unimproved Access 




102 ±11.30 


165 ±19.59 


0.004 



P-values are the result of 1-way ANOVA's. 



located significantly closer to unimproved access 
and further from hiding cover than expected on the 
basis of availability. 

Bedding sites had a significantly higher percent 
grass cover, and significantly less spruce, pine and 
fir in the canopy than control plots (Table 5). Elk 
selected bedding sites with significantly fewer trees 
per ha than available. There was no significant dif- 
ference between bedding sites and the control plots 
in terms of the distance to major and minor roads, 
and the distance to unimproved access (Table 5). 
There was no significant difference between the dis- 
tance from the bedding site to foraging areas and the 
distance from the control plots to foraging areas 
(Table 5). 



Discusion 

The results from multi-scale habitat studies are 
largely dependent on the selection of variables cho- 
sen to investigate (Pedlar et al. 1997). Erroneous 
conclusions may occur if important variables are 
selected at one scale and frivolous variables are 
selected at another (Pedlar et al. 1997). Selection of 
variables based on the findings of previous studies 
may alleviate the problem of variable selection. In 
our study we selected variables at each scale that 
represented characteristics of food, thermal cover 
and hiding cover; requisites that have been deemed 
important for the selection of habitat types and 
features by elk. Therefore we believe reasonable 
variables were investigated and should provide 



Table 4. Mean attributes at elk feeding (n=96) and control plots (n=96) in west-central Alberta during the 
winter of 1994- 1995 





Elk Feeding Sites 


Control plots 
Mean ± SE 






M±SE 


P Value^ 


Non-spatial Variables 








Canopy Closure (%) 


31.34 ±3.45 


63.72 ±3.55 


< 0.001 


Grass Cover (%) 


59.73 ± 3.37 


32.61 ±3.20 


< 0.001 


Herb Cover (%) 


19.00± 1.16 


19.85 ±1.01 


0.574 


Shrub Cover (%) 


15.31 ± 1.50 


26.25 ± 1.95 


< 0.001 


Spruce, Pine, & Fir in the canopy (%) 


28.17 ±4.20 


44.70 ± 4.06 


0.004 


Stem Density (# / ha X 1000) 


1.31 ±0.19 


3.24 ± 0.29 


<0.{X)1 


Tree Height (m) 


5.46 ± 0.72 


12.44 ±0.78 


< 0.001 


Spatial Variables 








Major Road 


1093 ±66.06 


1088 ±64.50 


0.829 


Minor Road 


601 ±70.05 


635 ± 72.24 


0.139 


Unimproved Access 


102 ± 11.30 


164± 12.34 


< 0.001 


Thermal Cover 


160± 14.35 


158 ± 18.25 


0.932 


Hiding Cover 


71 ±7.68 


48 ± 8.06 


0.024 



■^ P-values are the result from paired t-test. 



188 



The Canadian Field-Naturalist 



Vol. 116 



Table 5. Mean attributes at elk bedding (n=62) and control plots (n=62) in west-central Alberta during the 
winter of 1994-1995 





Elk Feeding Sites 


Control plots 
Mean ± SE 






M±SE 


P Value^ 


Non-spatial Variables 








Canopy Closure (%) 


64.68 ± 4.03 


71.74 ±4.01 


0.157 


Grass Cover (%) 


34.95 ±3.30 


25.35 ± 2.92 


0.016 


Herb Cover (%) 


16.85 ± 0.89 


17.71 ± 1.29 


0.56 


Shrub Cover (%) 


27.37 ±1.76 


28.26 ±2.13 


0.698 


Spruce, Pine, & Fir in the canopy (%) 


38.53 ± 5.07 


54.21 ±5.15 


0.023 


Stem Density(# / ha X 1 000) 


2.03 ±0.17 


3.17 ±0.27 


0.001 


Tree Height (m) 


13.38 ±0.87 


13.75 ±0.78 


0.701 


Spatial Variables 








Major Road 


1244 ±87.46 


1229 ± 84.34 


0.550 


Minor Road 


763 ± 69.90 


757 ± 72.24 


0.830 


Unimproved Access 


165 ± 19.59 


155 ±19.50 


0.684 


Forage Stands 


71±11.14 


69 ±13.63 


0.880 



P-values are results from paired t-test. 



meaningful insight into the habitat selection patterns 
of elk at multiple scales. 

During winter, vegetation is at its lowest quality 
and quantity and therefore elk should select areas 
containing the highest quantity of available forage. 
Selecting areas of high forage concentration would 
reduce energy expenditures associated with locomo- 
tion through snow and decreased temperatures (Geist 
1982). Previous studies have shown elk prefer to 
graze on grasses (site selection) when snow depths 
do not impede foraging (Nelson and Leege 1982; 
Morgantini 1988). The selection of grass/meadows 
as foraging sites at the stand level has been well doc- 
umented (Craighead et al. 1973; Morgantini and 
Hudson 1979; Irwin and Peek 1983; Jenkins and 
Wright 1986; Morgantini 1988). The proportion of a 
home range as forage or habitat diversity (e.g., patch 
size and density) may be variables that are represen- 
tative of a landscape scale for elk. Previous studies 
have indicated habitat selection by elk at the land- 
scape level. Irwin and Peek (1983) concluded that 
forage density strongly influenced the size and loca- 
tion of elk home ranges in Idaho. In dry shrub steppe 
environments, larger home ranges may compensate 
for low forage availability (McCorquodale et al. 
1989). Black et al. (1976) proposed that optimal elk 
habitat consists of 40% of the land base as cover and 
60% as food in the proper size and spatial arrange- 
ment. As an ecotone species, elk select heteroge- 
neous areas for winter home ranges providing the 
opportunity to feed in grass meadows while taking 
advantage of the adjacent areas for cover (Geist 
1982). 

The present study confirms previous reports on 
selection of forage characteristics by elk at different 
scales. At the site scale elk selected open areas with 
high grass cover for feeding when compared to bed- 
ding locations and available habitat. At the stand 



level elk selected grass/meadows due to their con- 
centration of grasses while utilizing all other habitat 
types in proportion to their availability. The results 
of the landscape level analysis are less conclusive. If 
habitat diversity, which is represented by patch size 
and density, is representative of a need for a hetero- 
geneous environment to meet different life requi- 
sites, then elk did exhibit landscape level selection. 
Though not significant in comparison to available 
home ranges, elk home ranges showed little variation 
in the proportion classified as forage even though 
there was a wide range of home range sizes. This 
suggests that forage maybe an important factor in 
determining home range size and location in our 
study. 

Closed canopies can reduce thermoregulatory 
costs, reduce snow depths, and provide security for 
elk (Peek et al. 1982; Skovlin 1982; Peek and Scott 
1985). Thermal cover is provided by stands that pro- 
tect against extreme temperatures, whereas hiding 
cover provides a sense of security from predators 
and human disturbance. Much debate has centered 
around whether elk select closed stands for their 
thermal or hiding cover value (Geist 1982; Cook et 
al. 1998). Beall (1976) concluded that elk selected 
closed stands to reduce heat loss, whereas Peek and 
Scott (1985) believed that thermal cover would have 
an insignificant effect on energy requirements. They 
concluded that security cover was important in the 
presence of human disturbance (Peek and Scott 
1985). In western Alberta, Morgantini and Hudson 
(1979) concluded that habitat selection by elk in 
winter was not a response to the thermal environ- 
ment, but governed by human disturbance. 

Our results suggest that elk did not select for ther- 
mal cover requirements at any scale. At the land- 
scape level the proportion of the home range classi- 
fied as thermal cover was comparable to that of the 






2002 



Jones and Hudson: Winter Habitat Selection by American Elk 



189 



AHR, while at the stand level those habitat types that 
would represent thermal cover (e.g., closed mature 
or closed old-growth) were used in proportion to 
their availability. At the site scale elk selected bed- 
ding sites whose structural characteristics would be 
considered of lower thermal cover quality than the 
available habitat. 

The need for hiding cover by elk is well docu- 
mented (Hershey and Leege 1976; Morgantini and 
Hudson 1979; Peek and Scott 1985). Most studies 
have described the need for hiding cover in regards 
to human disturbance and, in particular, roads. The 
avoidance of habitat near open roads by elk has been 
extensively documented (Hershey and Leege 1976; 
Lyon 1979; Rost and Bailey 1979). Most studies 
have established avoidances of roads in terms of a 
distance, but Lyon (1983) determined avoidance or 
loss of habitat in terms of road density. Habitat 
effectiveness decreases dramatically as the density of 
roads increases such that an area with a road density 
greater than 0.62 km/km^ is reduced by 40% (Lyon 
1983). 

The results of selection by elk at multiple scales 
in terms of security requirements are less conclusive 
than those found for forage and thermal cover 
requirements. At the landscape level there was not a 
significant difference in the proportion of hiding 
cover comprising either the elk home ranges or 
AHR. This is not surprising as the study occurred in 
an area that is predominantly forested. We would 
argue that selection at the landscape level did occur 
in terms of hiding cover requirements based on the 
results of the road density analysis. Road densities 
for the elk home ranges were significantly less than 
those of the AHR. In addition the densities are below 
the 0.62 km/km^ density estabhshed by Lyon (1983). 

Previous studies of habitat selection have been 
conducted at a single scale, with results being 
extrapolated to higher or lower levels (Pedlar et al. 
1997). The appropriateness of extrapolating results 
to other levels has not been examined. Our results 
involving selection by elk of forage characteristics at 
the three levels would support the extrapolation of 
results to higher or lower levels. Based on our analy- 
sis, conclusive results for habitat selection were 
observed at the site and stand scale with inconclusive 
results at the landscape scale. This suggests that 
habitat selection, in terms of forage requirements, 
are bottom up as opposed to top down. There was no 
selection detected at any level by elk to meet the 
needs of the life requisite of thermal cover, therefore 
extrapolation to other levels would be appropriate 
(though unnecessary). The appropriateness of extrap- 
olating the results of the hiding cover analysis is less 
conclusive. Selection of habitat characteristics to 
meet the requirement of hiding cover was not evi- 
dent at the stand or site level, but may have been 
masked by the selection at the landscape level. 



During different times of the year and under 
increased levels of human disturbance selection for 
hiding cover may be more evident at the stand and 
site level. 

We acknowledge that our conclusions are based on 
the analysis of habitat selection data from one winter 
with lower than average precipitation levels. 
Because of this we did not examine the influence that 
snow depth had on habitat selection. It was also 
beyond the scope of our study to examine other 
factors (e.g., predation, inter and intra- specific com- 
petition) that can influence habitat selection. To over- 
come these limitations, further investigation is war- 
ranted to determine how these other factors change 
habitat selection patterns and our conclusions. The 
value of our study is to show resource managers the 
need to examine habitat selection patterns across 
more than one level to better manage elk. 

Acknowledgments 

We gratefully acknowledge the financial support 
of the Foothills Model Forest, without which this 
project could never have been completed. In addition 
we would hke to thank the Hinton Fish and Game 
Association and the Rocky Mountain Elk Foundation 
for their contributions. Equipment and logistic sup- 
port was provided by Weldwood of Canada Inc. 
(Hinton Division), Alberta Environmental Protec- 
tion, Fish and Wildlife Division (Edson), NOVA 
Gas Corporation, and The Environmental Training 
Centre. We thank F. Nelson of the Athabasca Ranch 
for permission to travel across her property. To D. 
Jewison, N. Gataiant, S. Kathnelson, W. Jones, and 
the many volunteers who helped in the collaring of 
elk and the collection of field data. Special thanks to 
M. Todd and D. Farr for their timely advice and 
guidance. 

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Received 31 March 2001 
Accepted 20 May 2002 



Notable Vascular Plants from Alaska in Wrangell-St. Elias 
National Park and Preserve with Comments on the Floristics 

Mary B. Cooki and Carl A. Roland^ 

•Wrangell-St. Elias National Park and Preserve, P.O. Box 439 Copper Center, Alaska 99573, USA 
2Denali National Park and Preserve, P.O. Box 9, Denali Park, Alaska 99755, USA 

Cook, Mary B., and Carl A. Roland. 2002. Notable vascular paints from Alaska in Wrangell-St. Elias National Park and 
Preserve with comments on the floristics. Canadian Field-Naturalist 1 16(2): 192-304. 

An inventory of the vascular flora north of the Bagley Icefield in Wrangell-St. Elias National Park and Preserve, Alaska, 
U.S.A., was conducted from 1994 to 1997. The objectives of the inventory were to assess the genetic diversity of the 
region, identify rare taxa and areas of phytogeographic interest and to assist park managers with planning and environmen- 
tal compliance. The purpose of this paper is to present annotations for 212 of the most notable taxa with their Alaska- 
Yukon and Park distribution maps and to summarize the floristics of the Park. 832 species (887 taxa) were documented as 
occurring within the Park (approximately 54% of the Alaskan flora). Significant results from the inventory include: 
(1) nine additions to the flora of Alaska (Arabis calderi G. A. Mulligan, A. codyi G. A. Mulligan, A. drepanoloba Greene, 
A. lemmonii S. Wats., Carex petasata Dewey, Draba lonchocarpa Rydb. var. thompsonii (C. L. Hitchc.) Rollins, Festuca 
minutiflora Rydb., Najas flexilis (Willdenow) Rostkov. & Schmidt and Trichophorum pumilum (Vahl.) Schinz & Thell. 
var. rollandii (Fern.) Hulten); (2) 331 range extensions; (3) the addition of 40 Alaska Natural Heritage plant taxa with a 
state rank of three or less for a total of 69 rare plant taxa in the Park's flora, and (4) the addition of 214 species to the 
Park's flora. The composition of the flora in percentages, with the rare flora in parenthesis, is: 25.8 (8.3) circumpolar, 
10.8 (9.7) incompletely circumpolar, 32.2 (38.9) North American, 23.2 (11.1) amphiberingian, 0.9 (1.4) amphiatlantic and 
7.1 (26.4) Alaska- Yukon endemic. The Alaska- Yukon and amphiberingian endemic elements are dominant in the Alaska 
Range, the Wrangell Mountains and in the Tanana, White and Copper River basins whereas the Cordilleran endemic 
element is dominant in the St. Elias and Chugach Mountains and Chitina River basin. These trends correspond to our 
understanding of plant migration after the Pleistocene Epoch from refugia in the Upper Yukon Valley, the Alaska Range 
and Beringia, the northern part of the Park being closest to these migration corridors, whereas the St. Elias and Chugach 
Mountains are at the northern end of the North American cordilleran migration corridor. Endemics may also have evolved 
from relict populations in unglaciated areas within the Late Wisconsin ice sheet adjacent to Lake Ahtna, on coastal refugia 
and on exposed sites in the dry northern interior of the Park bordering the Tanana Valley and the southeastern edge of 
Beringia. The rare flora was distributed somewhat evenly through the mountain ranges of the Park, but unevenly through- 
out the basin regions of the Park where the Chitina River basin had two to four times the number of rare species found in 
the other basins. There was a trend for rare plants to occur in the alpine zone, above 1200 m elevation, in xeric sites, in the 
alpine-herb talus slope plant community, on southerly aspects and on slopes of 20-40 degrees. Although the Park is 
protected, infrastructure development, visitor use in the backcountry in communities most likely to harbor rare plants, and 
global changes to the ecosystems are increasing. This inventory enhanced our understanding of the genetic diversity, 
phytogeography and distribution of our flora, however, there are still large areas in the Park that have not been surveyed, 
particularly in the coastal region, and we lack sufficient knowledge about most rare species (distribution, life history and 
population ecology) to develop protection plans. 

Key Words: Vascular plants, flora, rare plants, floristics, biogeography, range extensions. National Park, Alaska. 

Wrangeli-St. Elias National Park and Preserve Elias region for many boreal-montane and arctic- 

encompasses a unique cross section of boreal and alpine taxa that would be expected to occur here, 

coastal ecosystems in south-central Alaska with Published floristic studies within the Park were limit- 

floristic influences from Beringia, the Yukon, the ed to a few geographic areas: Skolai Creek, Chitistone 

arctic and the Pacific Mountain systems (Figure 1). Pass and the Chitistone River (Murray 1968, 1971); 

There is a high diversity of plant communities in this Russell, Sheep and Guerin Glaciers (Murray 1971); 

region due in part to the large size of the Park (5.3 Chitistone, Skolai and Frederika Valleys (Scott 1968, 

million hectares), the three climatic zones it covers 1974); Long Glacier and Dadina River (Saltmarch 

(interior, montane and transitional) and the wide 1978) and Bonanza Ridge near Kennicott (Nordell 

variety of landforms and lithologies found within its and Schmitt 1977). Other collections were made prior 

boundaries. to the establishment of the Park, but these were pri- 

Eric Hulten, in his history of botanical exploration marily along the two roads into the Park. Collections 
in Alaska, identified the Wrangell-St. Elias region as made from 1984 to 1992 during Park vegetation stud- 
one of three areas in Alaska that was poorly known ies documented 121 range extensions, twenty-two 
(Hulten 1940). The distribution maps in his flora of new rare plant species and verified that large areas of 
Alaska (Hulten 1968) show a void in the Wrangell-St. the Park were still unsurveyed. 

192 



2002 



Cook and Roland: Notable Vascular Plants from Alaska 



193 









62''30'N — 




A 



Park Boundary 
Roads 



100 Km 



Figure L Map of study area showing major geographical features of Wrangell-St. Elias National Park and Preserve, 
Alaska. 



The Park was established in 1980 under the 
Alaska National Interest Lands Conservation Act 
(ANILCA, Public Law 96-487). An inventory of the 
plants occurring within the Park is necessary to ful- 
fill the following management objectives of the Park 
as stated in the Final Environmental Statement 
(Alaska Planning Group 1973): 

1. Ensure retention of the magnificent Wrangell- 
St. Elias landscapes and living systems in a 
natural state. 

2. To the extent possible, allow the natural fluctu- 
ations and equilibrium of self-regulating 
ecosystems to continue unimpeded. 

3. Develop and implement a viable research 
program to provide basic information required 
for effective Park management. 

4. Lay early emphasis on identification of espe- 
cially fragile areas through appropriate research. 

The Natural Resources Management Guideline for 
the National Park Service, NPS-77 (USDI 1991) 



states that, "A basic vegetation inventory is the first 
step in vegetation management", and the General 
Management Plan for the Park mandates that the 
presence and extent of endangered species of flora 
and fauna be determined (USDI 1986). Vegetation in 
the Park is subject to alteration from recreational and 
subsistence uses including mining, timber harvest, 
grazing, all-terrain vehicles, hiking, climbing and 
camping. Backcountry use is increasing steadily as 
the Park infrastructure is developed and as the public 
seeks to experience new premier wilderness areas. 
Identification of sensitive taxa, their habitats, and 
unique tloristic associations was deemed necessary 
by the National Park Service for the preparation oi' 
backcountry management plans, environmental 
assessment and compliance and for the monitoring 
of subsistence uses. 

We conducted an inventory of the vascular tlora 
of selected areas north of the Bagley icefield within 
the Park from 1994 to 1997. The objectives o{' the 



194 



The Canadian Field-Naturalist 



Vol. 116 



inventory were to: increase our understanding of the 
ecological history, genetic diversity and biogeogra- 
phy of the Park's flora; identify populations of 
rare taxa, unique floristic associations and areas of 
phytogeographic interest in order to protect the 
biodiversity of the natural systems within the Park, 
and to provide the structure for an ongoing assess- 
ment of the Park's flora. The purpose of this paper is 
to present the collection data for the most notable 
range extensions along with Park and Alaska- Yukon 
distribution maps and to summarize the floristics of 
the Park. 

Study Area 

Four large mountainous regions define the land- 
scape of the Park: the Wrangell Mountains, the St. 
Elias Mountains, the Chugach Mountains and the 
Alaska Range. The Wrangell Mountains, a volcanic 
range with peaks reaching 4995 m, dominate the 
landscape in the center of the Park. The most com- 
mon bedrock in the Wrangell Mountains is Wrangell 
Lava, although the southern Wrangells have large 
exposures of Chitistone and Nizina limestone, 
Nikolai Greenstone and metamorphosed sedimentary 
rocks (MacKevett 1978; Richter 1976). The St. Elias 
Mountains straddle the border with the Yukon 
Territory and are even more precipitous and heavily 
glaciated than the Wrangell Mountains with peaks 
rising to 5489 m. The most common surface rock 
types in the St. EUas Mountains are Wrangell lava, 
Nikolai greenstone and marine sedimentary rocks of 
the Hasen Creek formation such as argillite, chert, 
conglomerate, limestone and shale (MacKevett 
1978). The Chugach Mountains are heavily glaciated 
due to their proximity to the Gulf of Alaska and 
extensive glaciation during the Pleistocene as indi- 
cated by the steep-sided U-shaped valleys, obvious 
trimlines, hanging glaciers and sparsely vegetated 
morainal features. The lithology of the Chugach 
Mountains is dominated by rocks of the Valdez 
group, a marine sedimentary unit composed primari- 
ly of graywacke, argillite, slate and phyllite 
(MacKevett 1978). North of the Wrangell Mountains 
lies the eastern arc of the Alaska Range which lies 
south of the Denali Fault and is represented within 
the Park by the Mentasta and Nutzotin Mountains. 
The lithology of the eastern Alaska Range is largely 
composed of marine sedimentary rocks with expo- 
sures of Nikolai Greenstone and thin-bedded 
hmestone (Richter 1976). 

The Copper, Chitina, White and Tanana (includ- 
ing the Nabesna and Chisana Rivers) are major inte- 
rior river drainage systems found within the Park. 
These drainage basins are mantled with quaternary 
surficial deposits of diverse origins. Surficial 
deposits include drift from the Wisconsin and older 
glaciations, Eolian deposits, Holocene alluvium and 
lacustrine sediments. The principal depositional fea- 
ture in the Copper River basin was the presence of 



the huge pro-glacial Lake Ahtna, which formed 
behind an ice dam at the confluence of the Copper 
and Chitina Rivers during the Pleistocene Epoch. 
Lake Ahtna occupied more than 520 000 hectares of 
the Copper River basin to a maximum elevation of 
800 m on the flanks of the Wrangell Mountains 
(Ferrians 1989). In contrast to the large expanses of 
open, low elevation terrain present in the Copper 
River basin, the upper Nabesna and Chisana Rivers 
occupy relatively narrow valleys. The valley floors 
of these braided rivers lie within the active flood- 
plain of the streams. The surficial deposits in the 
large, open valleys of the Chitina and White Rivers 
are a mix of alluvium on the floodplains and glacial 
outwash and drift from the Wisconsin glaciation 
on surfaces that have not been reworked by fluvial 
processes. Large areas in the White River valley are 
blanketed with volcanic ash from the eruptions of 
Mt. Churchill that occurred 1900 and 1250 years 
ago. 

Precipitation in the Park ranges from a yearly 
average of 338 cm at Yakutat, located in the mar- 
itime climatic zone, to 20 cm at Slana in the interior 
climatic zone. Temperatures on the coast are mild 
ranging from a mean daily high of 15°C to a mean 
daily low of -9°C whereas temperatures at Slana 
range from a mean daily high of 20°C to a mean 
daily low of -25°C. 

Five ecoregions and three transitional areas are 
found within the Park (Gallant et. al. 1995). These 
are: (1) the coastal Western Hemlock-Sitka Spruce 
forest ecoregion which is found on the Malaspina 
Forelands between Icy and Yakutat Bays; (2) the 
Pacific Coastal mountain ecoregion which includes 
the Chugach Mountains, southern St. -Elias 
Mountains and the Bagley Icefield; (3) the Wrangell 
Mountain ecoregion, demarcated by the Wrangellia 
Terrane; (4) the Copper Plateau ecoregion, the area 
of pro-glacial Lake Ahtna; (5) the Interior Highlands 
ecoregion, delineated by the Yukon-Tanana Terrane 
in the northeast corner of the Park; (6) the Alaska 
Range transition ecoregion encompassing the 
Nutzotin and Mentasta Mountains; (7) the interior 
bottomlands transition ecoregion, the region north of 
the Nutzotin Mountains in the Tanana Lowlands, 
and (8) the Duke Depression transitional ecoregion, 
the area encompassing the White River along the 
Duke River fault. 

Most of the Park was covered by the Cordilleran 
ice sheet or by the large pro-glacial Lake Ahtna 
during the peak of the Wisconsin glaciation (Clague 
1992; Hamilton 1994; Hamilton and Thorson 1983). 
The major river valleys became ice-free and Lake 
Ahtna drained with the retreat of the glacier occupy- 
ing the mouth of the Copper River canyon by 9.4 ka 
(Denton 1974; Hamilton 1994). There may have 
been unglaciated areas within the Late Wisconsin ice 
sheet adjacent to Lake Ahtna (Hamilton and Thorson 
1983), on coastal refugia and on exposed sites in the 



I 



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Cook and Roland: Notable Vascular Plants from Alaska 



195 



dry northern interior of tiie Park bordering the 
Tanana Valley and the southeastern edge of Beringia 
(Hopkins 1967). 

Previous Botanical Collections 

Eight significant plant collections were made 
within the Park prior to its establishment in 1980. 
WiUiam L. Poto collected 175 specimens primarily 
along the Mt. Drum Trail in the central Wrangell 
Mountains in 1902 as a member of the United States 
Geological Survey (USGS) Mt. Wrangell and 
Central Copper River Region Exploration Expe- 
dition (Poto 1902*; Mendenhall and Schrader 1903; 
Mendenhall 1905; Hulten 1940). Frank Charles 
Schrader and G. H. Hartman collected in the north- 
em Wrangell Mountains between the Nabesna and 
Copper Rivers the same year as Poto as part of this 
expedition. The specimens of Poto, Schrader and 
Hartman are at the U.S. National Herbarium. David 
W. Eaton, DeLorme D. Caimes and H. F. Lambart 
collected along the Yukon border, in the Park in the 
Wrangell and St. Ehas Mountains from 1909-1913 
as part of the Yukon- Alaska International Boundary 
Survey (Caimes 1911, 1914; Macoun 1914; Hulten 
1941-1950; Green 1982). The collections of Eaton 
are at the U.S. National Herbarium, those of Caimes 
and Lambart are at Canadian Museum of Nature in 
Ottawa. Hamilton M. Laing, a biologist with the 
Department of Mines of British Columbia who was 
a member of the Mt. Logan expedition in 1925, 
collected 243 specimens at the head of the Chitina 
River primarily in the St. Elias and Chugach Ranges. 
His collection was classified by A. E. Porsild and is 
housed at the Canadian Museum of Nature in Ottawa 
(Porsild 1939; Lambart 1926a, 1926b; Hulten 
1941-1950). 

David F. and Barbara Murray collected at May 
Creek, Nizina, Chitistone Pass, Skolai Pass, Guerin 
Glacier, Russell Glacier and Sheep Glacier in the 
Wrangell-St. Elias Mountains from 1966 to 1981 
(Murray 1968, 1971). Their collections are at the 
University of Alaska Fairbanks Museum and at the 
Canadian Museum of Nature in Ottawa. Richard W. 
Scott collected at Frederika Glacier, Chitistone Pass 
and Snag Glacier in the Wrangell Mountains in 1967 
and 1968 while conducting an ecological phytogeo- 
graphical study of the southeastern Wrangell 
Mountain flora (Scott 1968, 1974). Scott's collection 
is at the University of Alaska Fairbanks Museum and 
at the University of Michigan. Olle Nordell and Alf 
Schmitt collected 207 specimens at Kennicott and 
Bonanza Ridge in the Wrangell Mountains in 1976 
(Nordell and Schmitt 1977). Their specimens are at 
Lund and the University of Alaska Fairbanks 
Museum. Ransom Saltmarch collected 76 specimens 
in 1978 on the slopes of Mt. Wrangell while con- 

*See Documents Cited section. 



ducting research for a novel (Saltmarch 1978). His 
specimens were reviewed by D. F. Murray (ALA). 

An additional 36 individuals collected primarily 
along the two roads into the Park or on the roads 
adjacent to the Park. Among these collectors are: 
Edwin F. Glenn (1899), Martin Woodlock Gorman 
(1898 and 1899), E. L. Blaschke (1820), Arthur 
James Collier (1902), Adolphus Washington (1905), 
I. E. Diehl (1908), Walter Harrison Evans (1909), 
Bayne-Beauchamp Expedition (1932), William 
Albert Setchell (1932), Frits Warmolt Went (1934), 
Jacob Peter Anderson (1935 and 1944), Elisabeth 
Kol (1936), H. M. Raup (1944), Artheme-Antoine 
Dutilly and Emest LePage (1945-1947), Eric Hulten 
(1961), M. Sharrock (1962), George Argus (1967), 
R. Pegau (1968, 1970), and Leslie Viereck 
(1957-1980). The only collections made along the 
Park's coast were eight made by Frederick Funston 
in 1892 near Esker Stream and Manby Point on the 
Malaspina Forelands, five made by the Harriman 
Expedition at the Hubbard Glacier in 1899 and 
numerous Salix specimens collected by George 
Argus near Esker Stream (Coville and Funston 1896; 
Hulten 1941-1950; Argus 1967). Park staff collected 
1145 specimens at 215 unique localities throughout 
the Park from 1982 to 1994 during research and 
resource management studies. 

Methods 

We conducted a reconnaissance inventory north of 
the Bagley Icefield from 1994-1997. Site selection 
focused on surveying a range of plant communities, 
lithologies (such as calcareous, ultramafic, basalt, 
ash and greenstone) and landforms (such as south- 
facing bluffs, sand dunes, warm springs and flood- 
plains). We also focused site selection on azonal 
communities (scree slopes, wetlands and aquatics) 
and those areas where we predicted rare and endemic 
species would occur. In all, 239 sites were surveyed 
and approximately 7000 specimens were collected. 

Specimens were critically examined by the 
authors, Alan Battan, David F. Murray and Carolyn 
Parker at the University of Alaska Museum and the 
following specialists: George W. Argus (CAN) — 
Salix; Reidar Elven (O) — Draha lactea and 
Cerastium regelii; Mark Egger (WTU) — Castilleja: 
Barbara Ertter (UCB) — Potentilla diversifolia and 
P. drummondii: Donald Farrar (ISC) — Botrychiwn: 
Signe Frederickson (C) — Festiica\ G. A. Mulligan 
(DAO) — Arahis and Draba; David F. Murray — 
Papaver and Carex section Atratae; and Marcia 
Waterway (MTMG) — Carex laxa. 

We developed a set of relational databases in 
which the biogeography, taxonomy and rarity of 
each species was recorded and site characteristics for 
each rare plant collection was dociinicnlcd and 
evaluated. Distribution maps were prepared for 
selected taxa by assembling the following locality 



196 



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Vol. 116 



data into a Geographic Information System: all 
collections from this inventory and previous Park 
collections, specimen records from the University of 
Alaska Fairbanks Museum Plant Documentation 
Center and published stations from regional floras 
and monographs (Aiken and Darbyshire 1990; Argus 
1973; Argus 2000; Bayer 1993; Cody 1996; Cody et. 
al. 1998; Hulten 1941-1950; Hulten 1968; Porsild 
and Cody 1980). The distribution maps are at two 
scales, one encompassing all of Alaska and the 
Yukon Territory including adjacent parts of Canada 
and one focused on the Park. Nomenclature follows 
Cody (1996) for scientific names and Kartesz and 
Meacham (1999*) for common names. 

Results and Discussion 

Taxonomic Composition 

There are 832 species documented by vouchers 
within the Park, with a total of 887 taxa including 
intraspecific taxa. The flora of the Yukon Territory 
has approximately 1199 species, the flora of Alaska 
has approximately 1535 species and the two regions 
combined have approximately 1654 species with 
1202 species in common (Kartesz and Meacham 
1999). The number of species in the Park's flora is 
therefore approximately 54% of the Alaskan flora 
and 69% of the Yukon Territory flora. For compari- 
son with other protected areas, Denali National Park 
and Preserve in Alaska reported 684 taxa in 2000 
and Kluane National Park in the Yukon Territory has 
recorded 704 taxa (656 species) (Caswell 2002*). 

The flora of the Park is distributed among 81 
families and 258 genera. Eight of the species are not 
native to Alaska. The most important plant families 
within the flora are the Cyperaceae with 100 species 
(104 taxa), the Poaceae with 69 species (84 taxa), 
the Asteraceae with 68 species (75 taxa) and the 
Brassicaceae with 66 species (71 taxa). 

Phytogeography 

The Wrangell-St. Elias region is located within 
the distribution patterns of the major North 
American floristic elements, hence the plant species 
here have diverse origins and histories. They include 
species endemic to the area, broadly distributed 
boreal forest plants, arctic-alpine species, endemics 
of the western mountains and amphiberingian taxa 
of Asian origin. We recognize the following floristic 
elements within the vascular flora of the Park: 

I. Circumpolar species — 25.8% of the flora (226 species). 
This group includes those broadly distributed species that 
occur on all circumpolar continents including both Asiatic 
and European regions of Eurasia, Greenland and North 
America. We further divide this group of plants into boreal 
species primarily found in lowland and montane habitats and 
arctic/alpine taxa that generally occur in treeless landscapes 
in the arctic or alpine regions. A third element of the circum- 
polar flora is a group of more wide-ranging species with 
broad ecological amplitudes that occur across both arctic and 



boreal zones. The ratio of these elements within the circum- 
polar flora and examples of taxa from each category are: 

(1) Arctic-alpine species (10.6%) — Cardamine bellidi- 
folia, Carex lachenallii, Draba fladnizensis, Erigeron 
humilis, Saxifraga oppositifolia and Silene acaulis. 

(2) Boreal-montane species (6.7%) — Carex tenuifolia, 
Linnaea borealis and Rosa acicularis. 

(3) Widespread species (8.5%) — Carex capillaris, 
Empetrum nigrum, Equisetum arvense and Poa 
glauca. 

II. Incompletely circumpolar species — 10.8% of the flora 
(94 species). Species in this group have distributions that 
are very similar to the circumpolar plants of boreal distri- 
bution, but are not known from either Greenland or Europe. 
Examples of incompletely circumpolar species include 
Calypso bulbosa, Carex limosa, Chamaedaphne calycula- 
ta, Equisetum fluviatile, and Moehringia lateriflora. 

III. North American species — 32.2% of the flora (282 
species). Taxa in this group have distributions that are gen- 
erally restricted to North America. Two additional groups 
of species with more narrow distributions can be identified 
within the North American element: cordilleran species, 
which occur in the western mountains, and Pacific coastal 
species, which are generally restricted to the coast ranges 
and the west coast of North America. 

(1) Arctic-alpine species (2.1%) — Erigeron composi- 
tus and Primula egaliksensis 

(2) Boreal-montane species (11.4%) — Shepherdia 
canadensis and Viburnum edule 

(3) Cordilleran species (10.2%) — Luetkea pectinata 
and Salix commutata 

(4) Pacific coastal species (6.2%) — Lupinus nootka- 
tensis and Epilobium luteum 

(5) Widespread species (2.4%) — Solidago multiradiata 

IV. Amphiberingian species — 23.2% of the flora (203 
species). These are species that occur in North America 
and northern Asia but are not known from either Greenland 
or Europe; hence their center of distribution generally lies 
within Beringia. We separate the species within the 
amphiberingian biogeographic element into the following 
categories: 

(1) Arctic-alpine species (8.9%) — Artemisia arctica 
and Senecio atropurpureus 

(2) Boreal-montane species (4.3%) — Achillea sibirica 
and Boschniakia rossica 

(3) Pacific coastal species (5.0%) — Carex macro- 
cheata and Fritillaria camschatcensis 

(4) Widespread species (5.0%) — Parrya nudicaulis 
and Carex podocarpa 

V. Amphiatlantic species — 0.9% of the flora (8 species). 
Taxa in this group of species have distributions that include 
North America, Greenland and Europe but are not known 
from Asia. Carex nardina and Draba crassifolia are exam- 
ples of this element. 

VI. Alaska-Yukon endemic species — 7.1% of the flora 
(62 species). These are species that are restricted to Alaska 
and Yukon Territory, and sometimes extend into neighbor- 
ing regions of Northwest Territiories and British Columbia. 

(1) Arctic-alpine species (2.3%) — Thlaspi arcticum 
and Smelowskia borealis 

(2) Boreal-montane species (2.2%) — Salix setchel- 
liana and Artemisia alaskana 

(3) Cordilleran species (1.4%) — Synthyris borealis 
and Stellaria alaskana 



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Cook and Roland: Notable Vascular Plants from Alaska 



197 



(4) Pacific coastal species (L3%) — Castilleja una- 
laschensis and Salix stolonifera 

The composition of our flora is very similar to that 
described by George Douglas for the southwest 
Yukon (Douglas 1974). The northerly elements 
(circumpolar, incompletely circumpolar, North 
American arctic-alpine, amphiatlantic and amphi- 
beringian) comprise 64% of both floras whereas the 
southern elements (Cordilleran and Pacific Coast) 
contribute 16% to each flora. The North American 
boreal-montane group contributes 14% to the south- 
west Yukon flora and 11% to the Wrangell-St. Elias 
flora. The southwest Yukon flora had eight non- 
native species in 1974 as does the current Wrangell- 
St. Elias flora. Alaska- Yukon endemics comprise 7% 
of our flora and 5% (32 species) of the southwest 
Yukon flora reported by Douglas (1974). Lausi and 
Nimis (1985) documented 23 Alaska-Yukon 
endemics for the southern Yukon in a phytosociolog- 
ical study along the Alaska Highway. Porsild (1951) 
estimated that amphiberingian species comprise 
approximately 33% of the Alaska- Yukon flora. 
Douglas noted the southward decrease in this 
element for the southwest Yukon flora (17%). The 
slight increase in the contribution of this element to 
the Wrangell-St. Elias flora (23%) would support 
this gradient. 

We found the ratio of arctic and coastal elements of 
the southern Wrangell Mountain flora to be 
similar to that described by R. W. Scott in his phyto- 
geographical study of the southeast Wrangell 
Mountains (Scott 1974). Scott's analysis described a 
total of 291 species with 21% arctic, 7% coastal, 55% 
montane and 17% boreal. Our analysis resulted in 334 
species for the southern Wrangells with 39% arctic- 
alpine, 23% boreal-montane, 12% cordilleran, 6% 
Pacific coastal, 20% widespread and 0.3% introduced. 



Distribution of the Endemic Elements 
within the Park 

We evaluated the ratio of the endemic elements 
(Alaska- Yukon, amphiberingian, cordilleran and 
Pacific coastal) within each of the mountain and 
basin regions in the Park to determine if the composi- 
tional patterns are related to the ecological history of 
the region (Figures 2 and 3). The Alaska- Yukon and 
amphiberingian endemic floras are dominant in the 
Alaska Range (39% and 40% respectively), the 
Wrangell Mountains (28% and 42%) and in the 
Tanana (58% and 18%), White (39% and 52%) and 
Copper River basins (43% and 35%). This trend cor- 
responds to our understanding of plant migration 
after the Pleistocene Epoch from refugia in the Upper 
Yukon Valley, the Alaska Range and Beringia, the 
northern part of the Park being closest to these migra- 
tion corridors (Hulten 1937; Murray, et. al. 1983; 
Yurtsev 1963; Murray 1995). The Cordilleran 
endemic flora is dominant in the St. Ehas Mountains 
(45%), Chugach Mountains (37%) and in the Chitina 
River basin (38%) as would be expected since the St. 
Ehas and Chugach Mountains are at the northern end 
of the North American cordilleran. Several of our 
rarest species are cordilleran endemics reaching their 
northern extent in the Park {Arabis calderi G. A. 
Mulligan, A. lemmonii S. Wats, Carex petasata 
Dewey, and Festuca minutiflora Rydb). The Pacific 
coastal endemic element decreases notably from 
south to north in both the mountain and basin ranges. 
This trend may indiciate that migration north from 
the coast is impeded by the mountain ranges or by the 
availability of suitable habitat. The strong latitudinal 
gradient may also indicate that the Pacific coastal 
species represent a younger element in our flora 
when compared to the other three elements, which 
are more evenly distributed throughout the basin and 




D Alaska-Yukon ■ Amphiberingean ^Cordilleran ^ Pacific Coastal 






Alaska Range 
(n=:130) 




Wrangells St. Elias (n=53) Chugach 
(n=207) (n=152) 

Mountain Regions 

Figure 2. Ratio of endemic taxa within mountain regions of Wrangell-St. Elias National 
Park and Preserve, Alaska. 



198 



The Canadian Field-Naturalist- 



Vol. 116 



mountain ranges. Endemics may also have evolved 
from relict populations in unglaciated areas within 
the Late Wisconsin ice sheet adjacent to Lake Ahtna 
(Hamilton and Thorson 1983), on coastal refugia and 
on exposed sites in the dry northern interior of the 
Park bordering the Xanana Valley and the southeast- 
em edge of Beringia (Hopkins 1967). 

The Rare Flora 

Sixty-nine species in the Park's flora have an 
Alaska Natural Heritage state rank less than three 
and are treated as rare species by the National Park 
Service, 40 of these are new to the Park's flora. 
None of the rare species are considered threatened or 
endangered by the U.S. Fish and Wildlife Service. 
However, three species were listed as Species of 
Concern by the U.S. Fish and Wildlife Service in 
1995 {Cryptantha shackletteana L.C. Higgins, Carex 
laxa Wahlenb. and Taraxacum carneocoloratum 
Nels.). 

The most important families in the rare flora are 
the Brassicaceae with 17 species, the Cyperaceae 
with 13 species and the Caryophyllaceae with seven 
species. The three dominant classes for the distribu- 
tion of our rare flora by form of rarity (Rabinowitz 
1981) are: constantly sparse and geographically 
restricted in a specific habitat (30 species), locally 
abundant in a specific habitat but restricted geo- 
graphically (17 species) or constantly sparse in 
a specific habitat but occurring over a large range 
(12 species). 

Trends in the distribution of the rare flora by site 
characteristics are summarized by the following 
values. The number in parenthesis is the percent of 
total sites with that particular value for a site charac- 
teristic evaluated using 239 sites and 423 rare plant 
collections. 82% (37%) of all rare plant collections 
were made in the alpine zone; 78.5% (41.8%) were 
above 1200 m; 61% (37%) were in a xeric moisture 
class, and 57% (19%) were on alpine herb-talus 
slopes. 57% of all plant collections had a slope of 
20-40 degrees and 45% were collected on southerly 
aspects. The remaining aspect classes were distribut- 
ed approximately equally (north — 17%, east — 
19% and west— 19%). 

The rare flora appears to be distributed relatively 
evenly throughout the mountain ranges (> 1067 m) 
in the Park. The Alaska Range has 32 rare species, 
six of these being unique to the Alaska Range; the 
Wrangell Mountains have 33 species with three 
species unique to the Wrangell Mountains; the St. 
Elias Range has 19 rare species, two of which are 
unique to the St. Elias Range; and the Chugach 
Mountains (including the Granite Range) have 28 
rare species, eight of which are unique to the 
Chugach Mountains. The distribution of rare plants 
is uneven for the river basins in the Park. The 
Chitina River Basin has 32 rare species, six of which 
are unique to this basin. The Tanana River has 13 



rare species, two being unique; the Copper River 
basin has 13 rare species, two being unique, and the 
White River basin has eight rare species, with no 
species unique to that basin. 

The biogeographic composition of the rare flora 
with examples of species from each element follow. 
I. North American species — 38.9% (28 species). 

(1) Arctic-alpine (1 species) — Primula egaliksensis 

(2) Boreal-montane (8 species) — Agoseris glauca, 
Carex parryana, Eriophorum viridi-carinatum and 
Maianthemum stellatum 

(3) Cordilleran (19 species) — Arabis calderi, A. lem- 
monii, Carex petasata and Festuca minutiflora 

n. Alaska-Yukon endemics — 26.4% (19 species). 

(1) Arctic-alpine (6 species) — Thlaspi arcticum and 
Douglasia arctica 

(2) Boreal-montane (3 species) — Cryptantha shacklet- 
teana, Lupinus kuschei and Salix setchelliana 

(3) Cordilleran (6 species) — Arabis codyi and Taraxa- 
cum carneocoloratum 

III. Amphiberingean species — 11.1% (8 species) 

(1) Arctic-alpine (5 species) — Festuca lenensis, Stel- 
laria umbellata and Trisetum sibiricum 

(2) Boreal-montane (1 species) — Potamogeton subsi- 
biricus 

(3) Pacific coastal (2 species) — Rumex beringensis 
and Salix hookeriana 

IV. Incompletely circumpolar — 9.7% (7 species) 

( 1 ) Arctic-alpine ( 1 species) — Cryptogramma stelleri 

(2) Boreal-montane (4 species) — Tricophorum pum- 
ilum and Carex laxa 

(3) Widespread (2) — Ceratophyllum demersum and 
Myriophyllum verticillatum 

V. Circumpolar — 8.3% (6 species) 

(1) Arctic-alpine (5 species) — Colpodium vahlianum 
and Cerastium regelii 

(2) Boreal-montane (1 species) — Viola selkerkii 
VII. Amphiatlantic — 1.4% (1 species) — Najas flexilis 

The main differences between the biogeography of 
the total flora and the rare flora are the increase in the 
ratio of Alaska- Yukon endemics in the rare flora 
which is only 7% of the total flora, and the decrease 
in the ratio of circumpolar and amphiberingian ele- 
ments in the rare flora which is 37% and 24% of the 
total flora respectively. The ecological distribution of 
the rare flora is: cordilleran (38%), boreal-montane 
(25%), arctic-alpine (24%), Pacific coastal (6%), 
temperate disjunct (6%) and widespread (3%). The 
primary differences in the ecological distribution of 
the rare flora as compared to the total flora are the 
increase in the cordilleran element in the rare flora, 
which is only 12% of the total flora and the decreases 
in the Pacific coastal and widespread elements in the 
rare flora which is 12% and 20% of the total flora 
respectively. 

Recommendations 

This inventory has enhanced our understanding of 
the biodiversity and phytogeography of the region and 
has filled in the 'Wrangell Void' for many species not Aj 
known to occur in the Park or thought to be rare. " 



2002 



Cook and Roland: Notable Vascular Plants from Alaska 



199 




Xanana 
(n=22) 



White (n=31) 



Copper 
(n=37) 

River Basins 



Chitina 
(n=37) 



Gulf (n=24) 



Figure 3. Ratio of endemic taxa within river basins and Gulf of Alaska, Wrangell-St. Elias 
National Park and Preserve. 



However, there are many regions and communities 
that have yet to be surveyed or need further work. 
Some of these are: the Gulf of Alaska between Yaku- 
tat and Icy Bays, the southern St. Elias Mountains and 
Bagley Icefield, the Tanana lowlands, the White River 
basin, wetland and aquatic communities throughout 
the Park, nunataks, areas of high endemisim on the 
north side of the Park in the Mentasta, Nutzotin and 
northern Wrangell Mountains, high elevation plateaus 
in the Jacksina River drainage, and rare plant habitat 
throughout the Park. 

Over half of the Park (3.9 million hectares) is 
designated wilderness, the remaining area (1.4 mil- 
lion hectares) is theoretically protected. Aircraft, 
motorboats and cabins are allowed in wilderness in 
Alaska for subsistence purposes and aircraft conces- 
sions for climbing, hiking and backpacking are not 
restricted in the Park. Most of the increase in back- 
country use is in areas that we have found likely to 
harbor rare plants. The rate of global changes to 
ecosystems and plant communities is increasing as 
are infrastructure development and visitor use in the 
Park. Therefore, there is a need to survey areas 
where we anticipate use as well as the unsurveyed 
regions so that we can begin to assess changes to 
plant populations and communities. We also need 
information on the distribution, life history and 
population ecology of most rare plants so that 
protection plans may be developed. 

Notable Collections 

We documented nine additions to the flora of 
Alaska, 211 species new to the Park's flora, 145 
range extensions greater than 200 km from previous- 
ly documented localities and 186 range extensions 
greater than 60 km from previously documented 
localities. An annotated list in the phylogenetic order 



used by William J. Cody in the Flora of the Yukon 
Territory (1966) is preceded by a synoptic list of the 
most notable taxa by status. Alaska- Yukon and Park 
distribution maps for each species follow the annota- 
tions and are in the order of the annotations. 
Specimens are housed in the herbarium at Wrangell- 
St. Elias National Park and Preserve headquarters in 
Copper Center, Alaska unless otherwise indicated. 

Synoptic List by Status 
Taxa new to Alaska (9) 

Arabis calderi 

Arabis codyi 

Arabis drepanoloba 

Arabis lemmonii 

Carex petasata 

Draba lonchocarpa var. thompsonii 

Festuca minutiflora 

Najas flexilis 

Trichophorum pumilum var. rollandii 

Range Extensions (203) 

Agoseris glauca 

Agrostis mertensii 

Agrostis thurberiana 

Alopecurus aipinus 

Aphragmus eschscholtzianus 

Antennaria media 

Arabis media 

Arenaria capillaris 

Arenaria longipedunculata 

Arnica amplexicaulis ssp. prima 

Arnica latifolia 

Arnica mollis 

Artemisia hyperborea 

Aster aipinus ssp. vierfuipperi 

Aster boreal is 

Astragalus adsurgens ssp. viciifolius 

Astragalus eucosmus ssp. sealei 

Astragalus harringtonii 



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Astragalus nutzotinensis 

Astragalus williamsii 

Botrychium ascendens 

Botrychium lanceolatum 

Botrychium minganense 

Botrychium pinnatum 

Braya glabella ssp. glabella 

Braya glabella ssp. purpurascens 

Callitriche anceps 

Callitriche hermaphroditica 

Caltha leptosepala 

Carex adelostoma 

Carex albo-nigra 

Carex buxbaumii 

Carex chordorrhiza 

Carex crawfordii 

Carex eburnea 

Carex fdifolia 

Carex holostoma 

Carex interior 

Carex krausei 

Carex lasiocarpa ssp. americana 

Carex laxa 

Carex lenticularis var. dolia 

Carex leptalea 

Carex nardina 

Carex nigricans 

Carex obtusata 

Carex parryana 

Carex pauciflora 

Carex petricosa 

Carex phaeocephala 

Carex praticola 

Carex stylo sa 

Carex viridula 

Carex williamsii 

Cassiope mertensiana 

Castilleja chrymactis 

Castilleja elegans 

Castilleja yukonis 

Cerastium regelii 

Ceratophyllum demersum 

Chamaerhodos erecta ssp. nutallii 

Cicuta maculata var. angustifolia 

Cladothamnus pyrolaeflorus 

Claytonia tube rasa 

Collomia linearis 

Colpodium vahlianum 

Coptis trifolia 

Cryptantha shackletteana 

Cryptogramma crispa var. sitchensis 

Cryptogramma stelleri 

Cystopteris montana 

Danthonia intermedia 

Delphinium brachycentrum 

Deschampsia brevifolia 

Douglasia alaskana 

Douglasia arctica 

Douglasia gormanii 

Draba cinerea 

Draba crassifolia 

Draba corymbosa 

Draba densifolia 

Draba incerta 

Draba kananaskis 



Draba lactea 

Draba macounii 

Draba oligosperma 

Draba palanderiana 

Draba porsildii 

Draba ruaxes 

Draba stenoloba 

Draba stenopetala 

Epilobium lactiflorum 

Epilobium luteum 

Erigeron caespitosus 

Erigeron grandiflorus ssp. arcticus 

Eriophorum callitrix 

Eriophorum viridi-carinatum 

Erysimum pallasii 

Euphrasia mollis 

Eutrema edwardsii 

Fauria crista-galli 

Festuca brevissima 

Festuca lenensis 

Festuca richardsonii 

Festuca saximontana 

Gentiana douglasiana 

Gentiana platypetala 

Gentianella tenella 

Glyceria pulchella 

Gymnocarpium jessoense ssp. parvulum 

Hackelia deflexa 

Halimolobos mollis 

Hippuris montana 

Impatiens noli-tangere 

Isoetes echinospora 

Juncus filiformis 

Juncus mertensianus 

Juniperus horizontalis 

Kobresia sibirica 

Kobresia simpliciuscula 

Lesquerella arctica 

Ligusticum scoticum ssp. hultenii 

Lupinus kuschei 

Maianthemum stellatum 

Minuartia biflora 

Minuartia dawsonensis 

Minuartia strict a 

Mitella pentandra 

Montia bostockii 

Myriophyllum verticillatum 

Nymphaea tetragona ssp. leibergii 

Osmorhiza depauperata 

Oxytropis campestris ssp. jordalii 

Oxytropis huddelsonii 

Oxytropis scammaniana 

Papaver alboroseum 

Papaver radicatum ssp. kluanense 

Papaver walpolei 

Penstemon gormanii 

Phacelia mollis 

Phippsia algida 

Phlox hoodii 

Phlox richardsonii 

Phyllodoce glanduliflora 

Plantago eriopoda 

Poa hispidula 

Podagrostis aequivalis 

Polystichum lonchitis 



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Potamogeton friesii 

Potamogeton pectinatus 

Potamogeton pusillus var. pusillus 

Potamogeton praelongus 

Potamogeton subsibiricus 

Potamogeton zosteriformis 

Potentilla arguta ssp. convallaria 

Potentilla biflora 

Potentilla diversifolia 

Potentilla drummondii 

Potentilla litoralis 

Potentilla rubricaulis 

Primula cuneifolia ssp. saxifragifolia 

Primula egaliksensis 

Puccinellia deschampsioides 

Puccinellia interior 

Ranunculus gelidus ssp. grayi 

Ranunculus pacificus 

Ranunculus pedatifidus ssp. affinis 

Ranunculus sulphureus var. sulphurus 

Ranunculus trichophyllus var. eradicatus 

Rumex acestosa ssp. alpestris 

Rumex beringensis 

Sagina saginoides 

Salix commutata 

Salix rotundifolia ssp. dodgeana 

Salix setchelliana 

Salix stolonifera 

Saussurea angustifolia ssp. yukonensis 

Saxifraga adscendens ssp. oregonensis 

Saxifraga bracteata 

Saxifraga foliolosa 

Selaginella sibirica 

Silene involucrata ssp. involucrata 

Silene menziesii 

Silene repens 

Silene uralensis ssp. uralensis 

Silene williamsii 

Smelowskia borealis 

Smelowskia calycina var. porsildii 

Sparganium minimum 

Stellaria alaskana 

Stellaria umbellata 

Subularia aquatica 

Swertia perennis 

Synthyris borealis 

Taraxacum carneocoloratum 

Taraxacum phymatocarpum 

Thlaspi arcticum 

Trisetum sibiricum ssp. litorale 

Vahlodea atropurpurea 

Veronica serpyllifolia ssp. humifusa 

Viola adunca 

Viola biflora 

Viola selkirkii 

Alaska Natural Heritage Program State Rare Plants 
(State Rank < = 3) (69) 

Agoseris glauca 
Agrostis thurberiana 
Aphragmus eschscholtzianus 
Arabis calderi 
Arabis codyi 



Arabis drepanoloba 

Arabis lemmonii 

Arenaria longipedunculata 

Arnica mollis 

Astragalus harringtonii 

Botrychium ascendens 

Carex adelostoma 

Carex crawfordii 

Carex ebumea 

Carex holostoma 

Carex interior 

Carex laxa 

Carex lenticularis var. dolia 

Carex parryana 

Carex petasata 

Cerastium regelii 

Ceratophyllum demersum 

Colpodium vahlianum 

Cryptantha shackletteana 

Cryptogramma stelleri 

Cystopteris montana 

Douglasia alaskana 

Douglasia arctica 

Douglasia gormanii 

Draba densifolia 

Draba incerta 

Draba kananaskis 

Draba lonchocarpa var. thompsonii 

Draba porsildii 

Draba ruaxes 

Draba stenopetala 

Eriophorum viridi-carinatum 

Eiysimum pallasii var. pallasii 

Festuca lenensis 

Festuca minutiflora 

Glyceria pulchella 

Juniperus horizontalis 

Lupinus kuschei 

Maianthemum stellatum 

Minuartia biflora 

Montia bostockii 

Myriophyllum verticillatum 

Najas flexilis 

Oxytropis huddelsonii 

Papaver alboroseum 

Papaver walpolei 

Phacelia mollis 

Phlox hoodii 

Phlox sibirica ssp. richardsonii 

Potamogeton subsibiricus 

Potentilla drummondii 

Potentilla rubricaulis 

Ranunculus pacificus 

Rumex beringensis 

Salix setchelliana 

Saxifraga adscendens ssp. oregonensis 

Smelowskia calycina var. porsildii 

Stellaria alaskana 

Stellaria umbellata 

Taraxacum carneocoloratum 

Thlaspi arcticum 

Trichophonim pumilum \ ar. rollandii 

Trisetum sibiricum ssp. litorale 

Viola selkirkii 



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Species Accounts 

Selaginellaceae 

Selaginella sibirica (Milde) Hieron, Siberian Spike- 
Moss — Chugach Mountains: rare in tundra, 
Granitic Creek, 1768 m, 6r6.19' N 142°55.03' W, 
M. Cook 94220A, 8 July 1994. Nutzotin 
Mountains: scattered in rock crevices, Lick Ridge, 
1554 m, 62°28.44' N 142°15.06' W, M. Cook 
95118, 29 June 1995. Wrangell Mountains: scat- 
tered on SE-facing bluff, Nabesna River, 1106 m, 
62°15.05' N 142°54.68' W, M. Cook 96189, C. 
Roland 96-161, 19 June 1996; outcrop crevices, 
Nikolai Mine, 1695 m, 61°27' N 142°39' W, Batten 
& Barker 96-055, 24 July 1996; outcrop, Nikolai 
Ridge, 1372 m, 61°26.'5' N 142°43' W, Batten & 
Barker 96-157, 26 July 1996 (ALA); rock ledges. 
Crystalline Hills, 1585 m, 6r23.59' N 143°31.85' 
W, M. Cook 95276, 19 July 1995; in moss under 
open willow scrub, Fish Creek, 1067 m, 62° 16.92' N 
142°58.99' W, M. Cook 95312, 27 July 1995. 

This amphiberingean species occurs in dry, rocky 
situations throughout northern Alaska. The Granitic Creek 
locality extends its range 272 km to the southeast into the 
Chugach Mountains from a station in the Alaska Range 
reported by Hulten (1968). It was also collected at several 
intermediate localities in the Wrangell and Nutzotin 
Mountains. Map 1. 

ISOETACEAE 

Isoetes echinospora Durieu (/. muricata Durieu), 
Quillwort — Wrangell Mountains: abundant, 
uprooted and lining shoreline. Lake 2870, 0.2 km N 
of Tanada Creek, 875 m, 62°31.38' N 143'^ 28.12' 
W, C Roland 95-096, 28 June 1995; few clumped in 
muck, 0.6 m deep water. Fox Farm Lake, 727 m, 
62°19.98' N 144°50.02' W, M. Cook 95308, 25 July 
1995; abundant submerged in 30 cm deep water. 
Lake 2990', 3.6 km west of Copper Lake, 911 m, 
62°24.53' N 143°42.68' W, C. Roland 96-913, 
11 August 1996. 

This boreal North American aquatic species has been 
found at a few widely separated sites across central Alaska 
and the Yukon Territory. The specimens cited above extend 
its range 80 km southeast into the Wrangell Mountains 
from a station in the Alaska Range (Hulten 1968). Map 2. 

Ophioglossaceae 

Botrychium ascendens W. H. Wagner, Triangle- 
Lobe Moon wort — Nutzotin Mountains: few on 
S-facing scree slope. Gold Hill, 1434 m, M. Cook 
3520B, 18 July 1999. 

This North American species with a cordilleran distribu- 
tion was known from two localities in Alaska and one in 
the Yukon Territory (Cody 1994). It is rare in Alaska (G3? 
SI) and Cody (1994) suggested that it be added to the list 
of rare species for the Yukon Territory. The locality cited 
above extends its range 266 km to the southwest of a 
station in the Yukon Territory and 356 km northwest of a 
collection locality in the Yakutat Quad (Akwe Dunes, 
59°21.78' N 138°50.45' W, M.C. Stensvold 7304, 27 June 
1996 (ALA)). Map 3. 



Botrychium lanceolatum (Gmel.) Angstr., Lanceleaf 
Grapefern — Chugach Mountains: rare on S-fac- 
ing slopes. Juniper Island, 1346 m, 60''36.17' N 
142°14.96' W, C. Roland 94-208, 10 July 1994; 
openings in willow thicket. Granite Creek, 823 m, 
60°44' N 142°13' W, Batten & Barker 96-354B, 
30 July 1996 (ALA); open low shrub birch scrub. 
Granite Creek, 579 m, 60°44' N 142°32' W, Parker 
& Duffy 6718, 6 August 1996. Wrangell Moun- 
tains: earth-flow scar on steep S-facing slope, 
Nikolai Pass, 1372 m, 6r26.5' N 142°43' W, Batten 
& Barker 96-174B, 26 July 1996 (ALA); scattered in 
grassy gully. Long Glacier, 1399 m, 61°48.04' N 
144°6.27' W, M. Cook 96702, C. Roland 96-851, 
6 August 1996. 

This moonwort occurs in maritime-influenced boreal 
regions and scattered inland locations throughout the cir- 
cumpolar north. In Alaska, it occurs most frequently along 
the southern coast from the panhandle through the Aleutian 
Islands. Our collections extend its range 175 km northeast 
into the Wrangell and Chugach Mountains from a collec- 
tion in the Cordova Quad (60°58' N 145° W, C L. Parker 
1830B, 12 August 1986 (ALA)) and connect the 
distribution 202 km to the east in the Yukon Territory 
(Cody 1996). Map 4. 

Botrychium minganense Victorin, Mingan Moon- 
wort — Nutzotin Mountains: few in Dryas- 
graminoid-forb herbaceous stringer at base of scree 
slope, headwaters of Alder Creek, 1554 m, 62°28.44' 
N 142°15.06' W, M. Cook 95139, 30 June 1995; 
open shrub birch-willow scrub. Ptarmigan Lake, 
1128 m, 6r50.12' N 14^9.15' W, Dujfy & Barnes 
96-296, 8 August 1996; few on S-facing scree slope. 
Gold Hill, 1434 m, M. Cook 3520C, 18 July 1999. 

This moonwort occurs in scattered localities across bore- 
al North America and as far south as Arizona in the western 
Cordillera. It is known from only two other localities in 
Alaska, 405 km to the northwest in the Healy Quad 
(Chulitna River, 62°59.75' N 150°6.83' W, C. Roland 
3493B, 18 August 1998 (ALA)) and 308 km to the south- 
east in the Yakutat Quad (59° 19.35' N 138°40.14' W, M. 
Stensvold 7313, 16 July 1998 (ALA)). Map 5. 

Botrychium pinnatum H. St. John {B. boreale (E. 
Fries) Milde), Northeastern Moonwort — Chugach 
Mountains: scrub thicket. Granite Creek, 1067 m, 
6r5.46' N 142°54.05' W, J. Bolivar 84-172, 
1 August 1984; rare in disturbed mineral soil, upper 
Golchonda Creek, 1402 m, 61°2.89' N 143°24.42' 
W, M. Cook 3200, 28 July 1998; openings in willow 
thicket. Granite River, 823 m, 60^44' N 142°13' W, 
Batten & Barker 96-354C, 30 July 1996. Nutzotin 
Mountains: few in Dr>'<3.y-graminoid tundra, head- 
waters of Alder Creek, 1554 m, 62°28.44' N 
142°15.06' W, M. Cook 95121, 29 June 1995. 
Wrangell Mountains: scattered in meadow, vie. 
Chitistone Falls, 1 177 m, 61°32.27' N 142''11.49' W, 
C. Roland 96-632, 22 July 1996; steep E-facing 
meadow, Nikolai Mine, 1695 m, 61°27' N 142°39' 
W, Batten & Barker 96-074B, 24 July 1996 (ALA); 



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1. Selaginella sibirica 




2. Isoetes echinospora 




3. Botrychium ascendens 




4. Botrychium lanceolatum 



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The Canadian Field-Naturalist 



Vol. 116 




5. Botrychium minganense 




6. Botrychium pinnatum 




7. Cryptogramma crispa var. sitchensis 




8. Cryptogramma stelleri 



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9. Cystopteris montana 




10. Gymnocarpium jessoense ssp. parvulum 




11. Polystichum lonchitis 




12. Juniperus horizontalis 



206 



The Canadian Field-Naturalist 



Vol. 116 



earthflow scar on steep slope, Nikolai Pass, 1372 m, 
61°26.5' N 142°43' W, Batten & Barker 96-174A, 
26 July 1996; moist silt near stream, W slope of 
Chitistone Mountain, 1219 m, 6r28' N 142°33' W, 
Batten & Barker 96-146, 26 July 1996 (ALA); 
patchy in bare area, vie. Long Glacier, 1399 m, 
61°48.04' N 144°6.27' W, M. Cook 96701, 96703, 
96716, 8 August 1996; few in mesic forb herbaceous 
vegetation, vie. Grant Creek, 1250 m, 6n7.65' N 
143°56.34' W, M. Cook 96367, 7 July 1996. 

This amphiberingean species occurs in scattered locaH- 
ties throughout boreal areas of North America. It was 
considered rare in the Yukon Territory by Douglas et al. in 
1981. The collections cited above extend its range 324 km 
east into the Wrangell Mountains, 341 km to the northeast 
into the Nutzotin Mountains and 367 km east into the 
Chugach Mountains from a collection in the Anchorage 
Quad (6r50.0' N 149° 18.0' W, Parker & Murray 2097, 
10 August 1989 (ALA)). These new localities also connect 
the range to the west with the range 256 km to the east in 
the Yukon Territory. Map 6. 

Pteridaceae 

Cryptogramma crispa (L.) R. Br. var. sitchensis 
(Rupr.) C. Christens., Alaska Parsley Fern — 
Chugach Mountains: rubble slope. Granite Creek, 
579 m, 60°44' N 142°32' W, Parker & Duffy 6709, 
6 August 1996; scattered in moss, Lake Creek, 
975 m, 61°11.4' N 143°49.45' W, M Cook 96401, 
13 July 1996; scree, Hanagita Peak, 1186 m, 
61°4.9r N 143°38.86' W, M. Cook 96421, 14 July 
1996; occasional on lateral moraine, 12 Mile Basin, 
1271 m, 60°48.85' N 142°33.52' W, M. Cook 
96636, 30 July 1996; dry boulder area. Upper Tebay 
Lake, 610 m, 6ri 1.5' N 144°23.5' W, L. A. & E. G. 
Viereck 11083, 1 July 1996. St. Elias Mountains: 
closed tall alder scrub on glacial rubble, Atrevida 
Glacier, 59°57.06' N 139°48.5' W, B. Haller s.n., 19 
August 1987. 

This fern variety is endemic to Alaska, the Yukon 
Territory, northern British Columbia and the District of 
Mackenzie, Northwest Territories where it grows in moist 
talus and rock crevices. The specimens cited above extend its 
range 293 km northeast into the Chugach Mountains from a 
station near Seward (Hulten 1968) and connect the range 200 
km to the southeast at Yakutat (Hulten 1968). Map 7. 

Cryptogramma stelleri (S.G. Gmel.) Prantl., Fragile 
Rock-Brake — Nutzotin Mountains: head of 
Sheep Creek, 6 km above confluence with Chisana 
River, 1067 m, 62°7.95' N 14r54.9' W, J. Bolivar 
84-82, 26 June 1984; rare under boulders on SW- 
facing slope, Carl Creek, 1920 m, 62°3.52' N 
141°36.27' W, M. Cook 95006, 14 June 1994; rare in 
rock crevices. Garden Hills, 1311 m, 62°18.44' N 
14ril.55' W, M. Cook 94166, 25 June 1994. 
Wrangell Mountains: rare in rock crevices, Monte 
Cristo Creek, 975 m, 62° 1 3.66' N 142°56.35' W, C 
Roland 96-147, 96-149, 18 June 1996; rare under 
boulders, Jaegar Mesa, 1893 m, 62°15.9' N 
143°1.24' W, C. Roland 95-133A, 4 July 1995. 



This fern is circumpolar and widespread but rare 
throughout its distribution. It is considered rare in both 
Alaska (G5 S2S3) and the Yukon Territory (Douglas et al. 
1981). Our collections extend its range 312 km south into 
the Nutzotin and Wrangell Mountains from a collection in 
the Big Delta Quad (Brigadier Road, 64°40.75' N 
146°12.38' W, Duffy & Lipkin 95-654, 12 July 1995 
(ALA)). Map 8. 

Dryopteridaceae 

Cystopteris montana (Lam.) Bernh., Mountain 
Fragile Fern — Chugach Mountains: SE-facing 
bench. Lower Bremner River, 457 m, 61°3.39' N 
144°26.8' W, K.A. Teare 1809, 20 July 1984. 
Mentasta Mountains: scattered at forest margin 
toe of talus slope, Totschunda Creek, 1280 m, 
62°27.85' N 142M0.68' W, C. Roland 96-298, 
24 June 1996. Wrangell Mountains: limestone 
rocks in alder thicket. Fish Creek, 1067 m, 62° 16.92' 
N 142°58.99' W, C. Roland 95-293, 29 July 1995; 
scattered in moss-forb tundra on limestone outcrop, 
Lakina Glacier, 1219 m, 6r33.64' N 143°19.01' W, 
C. Roland 96-689, M. Cook 96534, 24 July 1996. 

This fern is circumboreal and is considered rare in 
Alaska (G4S3) where it occurs in montane meadows and 
thickets, often on calcareous substrates. It had been collect- 
ed in the Wrangell Mountains (Bonanza Ridge, 61°30' N 
142°51' W, Nordell & Schmitt 580, 1976 (ALA)). The new 
collections cited above extend its range 114 km north into 
the Mentasta Mountains and 100 km south into the 
Chugach Mountains. Map 9. 

Gymnocarpium jessoense (Koidz.) Koidz. ssp. parvu- 
lum Sarvela (G. robertianum (Hoffm.) Newm.), 
Limestone Oak-Fern — Nutzotin Mountains: rare 
on rock ledges under low birch/willow scrub, Garden 
Hills, 1311 m, 62°18.44' N 14ril.55' W, M. Cook 
94773, 25 June 1994. 

This species occurs in boreal areas throughout the cir- 
cumpolar region except Greenland. In Alaska and the 
Yukon it is generally restricted to the central Yukon River 
drainage except for a single locality near Anchorage report- 
ed by Hulten (1968). It is considered reire in the Yukon 
Territory (Douglas et al. 1981). The specimen cited above 
extends its range 180 km south into the Nutzotin Mountains 
from a station near Chicken in the Yukon-Tanana uplands 
(Hulten 1968). Map 10. 

Polystichum lonchitis (L.) Roth, Holly Fern — 
Chugach Mountains: rare on outcrops, E Fork" 
Little Bremner River, 1219 m, 6r4.53' N 144°3.6' 
W, L. A. & E. G. Viereck 11104, 1 July 1996; rare 
on E-facing rocks of ridge, vie. Spirit Mountain, 
1204 m, 61°21.09' N 144°32.4' W, C. Roland 96- 
624, 17 July 1996; scattered on S-facing talus slope, 
vie. 12-mile Creek, Granite Range, 1335 m, 
60°49.7r N 142°33.34' W, C. Roland 96-813, 30 
July 1996. Gulf of Alaska: moraine, Chaix Hills 
Icy Bay, 427 m, 60°7.52' N 141°8.r W, B. Halle 
s.n., 24 August 1987. Map 11. 

This fern is circumpolar with a boreal-montane distribu- 
tion and is considered rare in the Yukon Territory (Douglas 



1 



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Cook and Roland: Notable Vascular Plants from Alaska 



207 



et al. 1981). Our collections extend its range 275 km south- 
east into the southern St. Elias Mountains from a station 
near Thompson Pass (61°08' N 145° 44' W, C L. Parker 
2462, 23 July 1990 (ALA)) and connect the distribution 
110 km to the southeast at Yakutat (F. Funston 126, 
15 August 1892 (US), Hulten 1941). 

CUPRESSACEAE 

Juniperus horizontalis Moench, Creeping Savin — 
Chugach Mountains: S-facing bank of lake, vie. 
East Fork Kiagna River, Granite Range, 1487 m, 
60°59.5' N 142°1' W, Batten & Barker 96-202, 27 
July 1996. St. Elias Mountains: sandy hillside, 
Hubert's Landing, Chitina River, 671 m, 61°2.7' N 
14r38' W, D. Miquelle 84-38, 15 July 1984; allu- 
vial floodplain of Clear Stream, Chitina River, 
488 m, 6r5.82' N 141°57.92' W, Duffy^ & Barnes 
96-005, 8 August 1996. Wrangell Mountains: 
small patch in gully within steppe, Dadina River 
bluff. Copper River, 393 m, 61°51.49' N 145°0.72' 
W, C Roland 96-304, 1 July 1996; scattered on S- 
facing bluff in aspen woodland, Crystalline Hills, 
602 m, 61°23.12' N 143°36.16' W, M. Cook 3129, 
15 June 1998 (ALA). 

This species occurs in dry situations across boreal North 
America but is considered rare in Alaska (G5 S1S2) where 
it is known only from the Copper River drainage, the vicin- 
ity of Anchorage and a single locality in the Tanana River 
valley. It is common on bluffs along the Chitina and middle 
Copper River drainages where it is often found growing 
with Juniperus communis L. It had been collected in the 
Wrangell Mountains (Bonanza Ridge, 6r30' N 142°51' W, 
Nordell & Schmitt s.n., 1976 (LD & ALA), Nordell and 
Schmitt (1977) and at the head of the Chitina River (//. M. 
Laing 10, 11 (CAN), Hulten (1941) and Porsild (1939). Our 
collections extend its range 25 km north into the Copper 
River valley. Map 12. 

Sparganiaceae 

Sparganium minimum (Hartm.) E. Fries (5. natans 
L.), Arctic Burr-Reed — Chugach Mountains: 
pond margin. Middle Fork of the Bremner River, 
869 m, 60°55.05' N 143°43.86' W, Dujfy & Barnes 
96-114, 6 August 1996; mud of small dried pond. 
Tana River Flats, 442 m, 6n4.45' N 142°57.21' W, 
Duffy & Barnes 96-135, 1 August 1996; scattered in 
pond, upper Tebay Lake, 579 m, 61°! 1' N 144''24' 
W, Parker & Gracz 6767, 1 August 1996. 
Wrangell Mountains: shallow water, vie. Tanada 
Creek, 927 m, 62°30.34' N 143°25.44' W, Moran & 
Roland 95-18, 29 June 1995. 

This species is circumboreal and is considered rare in the 
Yukon Territory (Douglas et al. 1981). The specimens cited 
above extend its range 130 km south into the Wrangell 
Mountains from a station near Paxson (Hulten 1968) and 280 
km east into the Chugach Mountains from a collection in the 
Anchorage Quad (Otter Lake, 61°17.53' N 149°44.17' W, 
Duf}- & Tande 1020, 3 August 1994 (ALA)). Map 13. 

Potamogetonaceae 

Potamogeton friesii Rupr., Fries' Pondweed — 

Wrangell Mountains: abundant in small, rock- 



floored pond in 30-100 cm of water. Fish Creek, 
1067 m, 62°16.92' N 142°58.99' W, C Roland 95- 
2S6A, 29July 1995. 

This circumpolar species is widespread but with wide 
gaps. The specimen cited above extends its range 163 km 
northwest into the Wrangell Mountains from a station near 
Tonsina (Hulten 1968) Map 14. 

Potamogeton pectinatus L., Sago Pondweed — 
Copper River Basln: floating next to shore in pond. 
Old Edgerton Highway, km 2.5, 415 m, 61°49.3' N 
145°10.7' W, M. Cook 95372, 5 August 1995. 

This locality extends the range of this pondweed 106 km 
east into the Copper River basin from a station in the 
Anchorage Quad (McRoberts Creek, 61°33.0' N 148°55.0' 
W, Batten & Reed 80-1 19A, 1 July 1980 (ALA)). Map 15 

Potamogeton praelongus Wulf., White-Stemmed 
Pondweed — Wrangell Mountains: Scattered in 
shallow water around pond, vie. Orange Hill, 884 m, 
62°12.63' N 142°52.28' W, C. Roland 96-121B, 
17 June 1996; shore of shallow lake, vie. Monte 
Cristo Creek, 1021 m, 62°14.07' N 142°56.85' W, C. 
Roland 96-141, 18 June 1996; common in deep water 
of pond, vie. of Copper Lake, 911 m, 62°24.53' N 
143°42.68' W, C Roland 96-908, 11 August 1996; 
common in deep water of lake, vie. Lake 2910. 
Copper River, 823 m, 62°26.35' N 143°40.77' W, C. 
Roland 96-926, 12 August 1996. Chugach 
Mountains: fresh pondweed shallows, East Flowers 
Lake, 404 m, 61°5.5' N 142°34.2' W, Duffy & 
Barnes 96-205, 8 August 1996; 1 m deep water, 
Chokosna Lake, 619 m, 6r27.45' N 143°49' W, C 
Roland 94 -328 A, 10 August 1994; rooted in organic 
muck, Lakina Pond, 792 m, 61°29.49' N 143°25.6' 
W, C. Roland 96-677, 24 July 1996. 

This pondweed. incompletely circumpolar and 
widespread in its distribution, is rare in the Yukon Territory 
(Douglas et al. 1981). Our collections extend its range 199 
km south into the Chugach Mountains from a collection 
near Chistochina (62°30.0' N 144°50.0' W, G. Smith 2302, 
16 May 1954 (ALA)). Map 16. 

Potamogeton pusillus L. var. pusillus {P. berchtoldii 
Fieber), Small Pondweed — Chugach Mountains: 
mixed freshwater herbaceous, pond margin 1 m deep. 
Tana Flats, 442 m, 6ri4.45' N, 142°57.21' W, Duffy 
& Barnes 96-130B, 96-127, 96-122B, 1 August 1996: 
floating adjacent to shore. Old Edgerton pond, 
415 m, 6r49.3' N 145°10.7' W, M. Cook 95374, 5 
August 1995; common in shallow water, Happel 
Slough, 180 m, 6r2.13' N 144°29.43' W, C Roland 
96-618, 17 July 1996. Wrangell Mountains: 
beaver pond E of Chititu Creek, 457 m, 61^22.16' N 
142°40.29' W, M. Duffy 91043, 8 July 1991; in 1 m 
deep water, Chokosna Lake. 619 m. 6r27.45' N 
143°49' w, C Roland 94-328C, 10 August 1994; 
common, submerged in shallow water, vie. Indian 
Creek, 655 m, 62"38.1' N 144''20.2' W, M. Cook 
95343, 1 August 1995; rare on bottom in 30 cm deep 
water. Lake 2990'. Copper River, 91 1 m. 62^^24.53' 



208 



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Vol. 116 



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2002 



Cook and Roland: Notable Vascular Plants from Alaska 



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2 0. Najas flexilis 



210 



The Canadian Field-Naturalist 



Vol. 116 



N 143°42.68' W, C. Roland 96-909, 11 August 
1996; common rooted in muddy bottom, Copper 
River pond 0.5 mi E of Lake 2910', 823 m, 62°26.35' 
N 143°40.77' W, C. Roland 96-925, 12 August 
1996; forming dense stands throughout bottom of 
lake. Fish Creek, 1067 m, 62°16.92' N 142°58.99' 
W, C. Roland 95-273, 28 July 1995, C. Roland 95- 
272, M. Cook 95321, 29 July 1995. 

This species is circumpolar with a widespread distribu- 
tion. The specimens cited above extend its range 168 km 
southeast into the Wrangell Mountains from a station near 
Paxson (HuUen 1968) and 99 km east into the Chugach 
Mountains from a station near Thompson Pass (Hulten 
1968). Map 17. 

Potamogeton subsibiricus Hagstr., Yenisei River 
Pond weed — Wrangell Mountains: rare in shal- 
low water, Lake 2990', upper Copper River Basin, 
911 m, 62°24.53' N 143°42.68' W, C. Roland 96- 
919, 11 August 1996. 

This amphiberingean species occurs in widely separated 
localities in boreal regions of Alaska and the Yukon. It is 
considered rare in Alaska (G3 S3) and the Yukon Territory 
(Douglas et al. 1981). The Copper River locality extends its 
range 87 km east into the Wrangell Mountains from a 
station along the Richardson Highway between Gakona and 
Paxson (Hulten 1968). Map 18. 

Potamogeton zosteriformis Fern., Flatstem 
Pondweed — Chugach Mountains: common just 
below surface next to shore, pond at mile 1.3 Old 
Edgerton Highway, 415 m, 61°49.3' N 145°10.7' W, 
M. Cook 95375, 5 August 1995. 

This North American boreal-montane species is of limt- 
ed distribution in the Yukon Territory (Cody 1994; Cody et 
al. 1998). The specimen cited above extends its range 188 
km to the northwest into the Copper River basin from a 
locality in the Anchorage Quad (Little Kiowa Lake, 
6I°15.5' N 149°39.38' W, Diijfy^ & Tande 1031, 4 August 
1994 (ALA)). Map 19. 

Najadaceae 

Najas flexilis (Willdenow) Rostkov. & Schmidt, 
Wavy Watemymph — Wrangell Mountains: shal- 
low water, Chokosna Lake, 619 m, 6r27.45' N 
143049' w, C. Roland 94-325, 10 August 1994. 

This species is new to the flora of Alaska and although it 
was collected in 1986 in the Fort Yukon Quad (Heglund 
Plot Lake 23, & Lake 522 near Preacher Creek, 66°01'N 
144° 42' W, P. Heglund 86-351, 86-363, 16 & 17 July 
1986 (ALA)), it was not reported. It is now known from a 
third locality in Alaska in the Anchorage Quad (Little 
Kiowa Lake, 61°15.48' N 149°39.38' W, Dwj^' & Tande 
1030, 4 August 1994). The Wrangell Mountains locality is 
243 km northeast of the Anchorage Quad collection site. 
This species has an amphiatlantic distribution, and is 
widespread across North America and Europe. It is now 
considered rare in Alaska (G5 S1S2). Map 20. 

POACEAE 

Agrostis mertensii Trin. {A. borealis Hartm.), Red 
Bentgrass — Chugach Mountains: scattered in 
meadow on S-facing slope, vie. Twelve-Mile Creek, 
1335 m, 60°49.71' N 142^33.34' W, C. Roland 96- 



808, 30 July 1996; scattered in subarctic lowland 
sedge wet meadow, upper Tebay Lake, 579 m, 
61°ir N 144°24' W, Parker & Gracz 6758, 
1 August 1996. Wrangell Mountains: scattered in 
lichen-heath tundra, plateau vie. Long Glacier, 1399 
m, 6r48.04' N 144°6.27' W, C. Roland 96-853, 
96-870, 6 August 1996. 

This species is amphiberingean with an arctic-alpine 
distribution. The specimens cited above extend the range 
95 km south into the Chugach Mountains from a collection 
on Bonanza Peak in the southern Wrangell Mountains 
(6r30' N 142°51' W, Nordell & Schmitt 490, 1976 (LD & 
ALA), Nordell & Schmitt s.n., 1978). The new localities 
connect the range 106 km to the west in the Anchorage 
Quad (Brilliant Glacier, 61°06' N 147°29' W, M. Diijfy 
s.n., 19 August 1993 (ALA)) with a locality 441 km to the 
southeast near Skagway (59°33.0' N 135°06.0' W, A.R. 
Batten 88-516, 30 August 1988 (ALA)). Map 21. 

Agrostis thurberiana A. S. Hitchc. (Podagrostis 
thurberiana (Hitchc.) Hult.), Thurber's Bentgrass — 
Chugach Mountains: herbaceous meadow, under 
Salix barclayi, vie. 12-mile Creek, 1335 m, 60°49.71' 
N 142°33.34' W, C. Roland 96-833, 1 July 1996. 
Gulf of Alaska: tall forb herbaceous slope, 
Amphitheather Knob, 427 m, 59°57' N 139°46' W, M. 
Cook 87-165, 19 August 1987. 

This western North American species of Bentgrass 
occurs in scatterd localities along the southern coast of 
Alaska where it is considered rare (G5 S2). Our collections 
connect its range 212 km to the southwest in the Seward 
Quad (Iron Mountain, 60°22.00' N 147°39.0' W, M. Dujfy- 
93-1095, 17 August 1993 (ALA)) with a station near 
Pelican 345 km to the southeast (Hulten 1968). Map 22. 

Alopecurus alpinus J.E. Smith (A. borealis Trin.), 
Mountain Foxtail — Nutzotin Mountains: abun- 
dant in mesic tundra on SW-facing slopes, Wiki 
Creek, 1411 m, 61^54.46' N 141^10.68' W, C. 
Roland 96-088, 10 June 1996; disturbed wet sedge 
meadow, Francis Creek, 1085 m, 61^51.93' N 
141°9.4' w, Duffy & Barnes 96-233, 8 August 1996. 
Wrangell Mountains: common in bare clay soil of 
frost boils, Jaegar Mesa, 1893 m, 62°15.9' N 
143°1.24' W, C. Roland 95-101, 30 June 1995; 
Lakes Plateau, 1890 m, 62M.4' N 143°23.5' W, 
M. Potkin 95-115, 95-142, 31 July 1995. 

The collections cited above extend the range of this 
circumpolar, arctic-alpine grass south 182 km into the 
Wrangell Mountains and 244 km into the Nutzotin 
Mountains from a station in the Tanacross Quad (W Fork 
of the Dennison, 63°50.00' N 142°15.0' W, Smith & 
Viereck 2395, 24 June 1954 (ALA)). Map 23. 

Colpodium vahlianum (Liebm.) Nevski (Puccinellia 
vahliana (Liebm.) Scribn. & Merr.), Vahl's Alkalai 
Grass — Wrangell Mountains: scattered in wet 
frost boils on limestone ridge. Cooper Pass, 1942 m, 
62°17.16' N 142°31.44' W, C. Roland 94-279, 24 
July 1994 (ALA). 

This arctic-alpine tundra grass is circumpolar and is 
considered rare in both Alaska (G4 S3) and the Yukon 



2002 



Cook and Roland: Notable Vascular Plants from Alaska 



211 




21. Agrostis mertensii 




22. Agrostis thurberiana 




23. Alopecurus alpinus 




24. Colpodium vahlianum 



212 



The Canadian Field-Naturalist 



Vol. 116 



Territory (Douglas et al. 1981). The collection cited above 
is 719 km south of the nearest locality on the arctic coast 
(Table Mountain Quad: Double Mt., 68°44.00' N 143°35.0' 
W, A.R. Batten 75-5 14a, 22 July 1975 (ALA)) and 318 km 
west of a station in the Yukon Territory near Haines 
Junction (Cody 1996). Map 24. 

Danthonia intermedia Vasey, Timber Wild Oat Grass 
— Chugach Mountains: abundant in turfy meadow 
on SE-facing slope, Granite Creek, 1829 m, 61°0.22' 
N 141° 51. r W, C. Roland 96-752, 28 July 1996; 
abundant among heath hummocks on S-facing slope, 
vie 12-mile Creek, 1335 m, 60°49.71' N 142°33.34' 
W, C Roland 96-811, 30 July 1996; occasional in 
open patches of birch scrub, Granite Creek, 884 m, 
60°44.63' N 142°6.05' W, M. Cook 96604, C. 
Roland 96-792, 29 July 1996; moist herbaceous bank, 
Short River Pond, 503 m, 61°5.35' N 141°56.3' W, 
Parker & Duffy 6687, 6 August 1996; scattered in 
subarctic lowland sedge wet meadow. Falls Creek, 
579 m, 61°11' N 144°24' W, Parker & Gracz 6760, 
1 August 1996; mixed mesic forb herbaceous mead- 
ow & dry S-facing slope. Middle Fork of the 
Bremner River, 869 m, 60°55.05' N 143°43.86' W, 
Dujfy & Barnes 96-087, 96-104, 8 August 1996. St. 
Elias Mountains: locally common in graminoid 
meadow, Blondie Ridge, 1951 m, 60°53.94' N 141° 
6.75' W, C Roland 95-171, 12 July 1995. Wrangell 
Mountains: dominant in dry graminoid herbaceous 
meadow, Cheshnina Plateau, 1399 m, 61°48.04' N 
144°6.27' W, M. Cook 96715, C Roland 96-848, 96- 
864, 5 August 1996; scattered in turfy meadow, 
Chitistone Falls, 1177 m, 61°32.27' N 142°11.49' W, 
C. Roland 96-635, 22 July 1996. 

This species occurs in boreal meadows across North 
America and in Kamchatka (Hulten 1968). In Alaska, it 
was reported from the Kenai Peninsula and near Anchorage 
by Hulten (1968). Our specimens extend its range northeast 
163 km into the Wrangell Mountains, 239 km into the St. 
Elias Mountains and 182 km into the Chugach Mountains 
from a collection near Cordova (60°30.93' N 145°25.04' 
W, C. Parker 1944, 15 August 1986 (ALA)) and connect 
the range 215 km to the east in the Yukon Territory near 
Haines Junction (Cody 1996). Map 25. 

Deschampsia brevifolia R. Br. (D. caespitosa ssp. 
brevifolia (R. Br.) Tzvelev), Tufted Hairgrass — 
Wrangell Mountains: patchy in wet graminoid 
tundra, vie. Sheep Glacier, 1478 m, 62°21.85' N 
144°23.73' W, M. Cook 94425, 30 July 1994 (ALA); 
Lakes Plateau, 1890 m, 62°4.4' N 143°23.5' W, M. 
Potkin 95-089, 29 July 1995; few on gravel bar. 
Copper River near Lake 2990', 911m, 62°24.53' N 
143M2.68' W, M. Cook 96737, 11 August 1996; 
scattered in openings between alders on gravel bar, 
vie. Batzulnetas, 640 m, 62°38.6' N 143°52.09' W, 
M. Cook 96749, 96752, 13 August 1996. 

The specimens cited above extend the range of this 
circumpolar, arctic-alpine species 291 km south into the 
Wrangell Mountains from a station in the Alaska Range 
(Huhen 1968) and connect the range 180 km to the east in 
the Yukon Territory (Cody 1996). Map 26. 



Festuca brevissima Jurtzev, Alaska Fescue — 1 
Chugach Mountains: scattered along ridge between 
Chakina River and Granitic Creek, 1768 m, 6r6.19' 
N 142°55.03' W, M. Cook 94227A, 1 July 1994. ^ 
Mentasta Mountains: bare soil of exposed creek 
cutbank. Lost Creek, 1722 m, 62°36.45' N 
143°12.03' W, Beck & Cook 95240, 4 July 1995. 
Nutzotin Mountains: scattered in orange rhyolite 
scree, plateau between Bryan and Willow Creeks, 
1829 m, 6r58.97' N 141°50.03' W, M. Cook 95052, 
18 June 1995; occasional in moss stringers on unsta- 
ble granitic rubble slope. Antler Creek, 1585 m, 
62°25.78' N 142°22.11' W, Cook & Beck 95103, 27 
June 1995; scattered in bare mineral soil on top of 
rock outcrop, alpine basin at head of Lick Creek, 
1554 m, 62°28.44' N 142°15.06' W, M. Cook 95125, 
29 June 1995; scattered on sheep trail along ridge, 
Carl Creek, 1554 m, 62°2.81' N 141°34.65' W, M. 
Cook 3146A, 1 July 1998 (ALA); scattered in moist 
gravel, Wiki Basin, 1585 m, 6^55.08' N 14ri2.07' 
W, M. Cook 3178, 15 July 1998 (ALA). Wrangell 
Mountains: common in dry gravel, NE slope of Mt. 
Drum, 1433 m, 62°8.83' N 144°30.18' W, C. Roland 
94-070, 11 June 1994; occasional on S-facing basalt 
outcrops, Cone Ridge, 2073 m, 62°8.21' N 
143''18.47' W, M. Cook 94388, 26 July 1994; occa- 
sional in unstable S-facing rubble and cinders. Cone 
Ridge, 2073 m, 62°8.21' N 143°18.47' W, C. Roland 
94-317, 27 July 1994; scattered in Cassiope- 
graminoid tundra on SW-facing slope, Chetaslina 
plateau, 1615 m, 61^56.51' N 144°25.93' W, M. 
Cook 94478, 16 August 1994; scattered in gravel on 
frost-boils, Jaegar Mesa, 1893 m, 62°15.9' N 
143°1.24' W, C. Roland 95-110, 95-114, 95-119, 95- 
131, 1 & 2 July 1995; scattered in gravel patches 
amongst Dryas tundra. Crystalline Hills, 1585 m, 
6r23.59' N 143°31.85' W, M. Cook 95266, 18 July 
1995; Lakes Plateau, 1890 m, 62°4.4' N 143°23.5' 
W, M. Potkin 95-090, 29 July 1995; moist bare min- 
eral soil of solifluction lobes. Black Mountain, 
1481 m, 62°20.85' N 143°44.9' W, M. Cook 95141, 
95159, 1 July 1995; scattered in bare gravel patches 
within Cassiope-QncdiCtoxxs heath, Boulder Lake, 
1036 m, 62°31.27' N 144^1 1.26' W, M. Cook 95357, 
8 August 1995, confirmed by Signe Fredericksen (C) 
1998; occasional in morainal deposits, valley 
between Ruddy Mountain and Mt. Drum, 1615 m, 
62''4.6r N 144°46.39' W, C. Roland 96-326, 2 July 
1996; N-facing scree, Nikolai Pass, 1280 m, 6^26' 
N 142°40' W, Batten & Barker 96-008, 23 July 
1996; bouldery dry tundra on south slope, Nikolai 
Pass, 1280 m, 6r26' N 142^40' W, Batten & Barker 
96-100, 24 July 1996 (ALA); scattered in bare, 
gravel-sized rhyolite scree on SE-facing slope, 
Kuskulana Pass, 1545 m, 61^33.72' N 143°39.7' W, 
C. Roland 96-709, 26 July 1996; occasional in bar- 
ren, gravelly sites on SW-facing slope, Snyder Peak, 
1524 m, 62°4.47' N 144°30.51' W, C. Roland 96- 
380, 5 My 1996. 



2002 



Cook and Roland: Notable Vascular Plants from Alaska 



213 




25. Danthonia intermedia 



26. Deschampsia brevifolia 





27. Festuca brevissima 



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28. Festuca lenensis 



214 



The Canadian Field-Naturalist 



Vol. 116 



This amphiberingean arctic-alpine species is rare in 
Alaska (G4 S3S4). The specimens cited above extend its 
range 265 km into the Chugach Mountains from a collec- 
tion in the Alaska Range (Independent Ridge, 63°40.0' N 
144°54.0' W, L.A. Spetzman s.n., 20 June 1957 (ALA)). It 
is fairly common at high elevations in the northern region 
of the Park, where it was collected at numerous localities in 
the Mentasta, Nutzotin and Wrangell mountains. Map 27. 

Festuca lenensis Drobov (F. ovina L. ssp. alaskensis 
Holmen), Tundra Fescue — Mentasta Mountains: 
common in gravelly sites on limestone ridge in dry 
Dryas octopetala tundra, Lost Creek, 1646 m, 
62°34.58' N 143^5.58' W, C. Roland 94-268B, 
23 July 1994. Nutzotin Mountains: scattered on 
unstable S-facing scree slope, Horsfeld Creek, 
1768 m, 62°2.88' N 141°13.18' W, M Cook 94152, C. 
Roland 94-129C, 24 June 1994; dense population in 
protected, turfy tundra, Klein Creek, 1747 m, 62°2.29' 
N 14ri9.88' W, C. Roland 95-034, 15 June 1995; 
scattered in fine, dry reddish soil with gravel, Wiki 
Creek, 1411 m, 61°54.46' N 14ri0.68' W, M. Cook 
96041, 96099, C. Roland 96-057, 11 June 1996 
(ALA); abundant in dry facies around outcrops, Rock 
Lake, 1119 m, 6r48.7' N 141°16.57' W, C. Roland 
96-038, 6 June 1996; moist, sandy grus with serai 
herbs on well-vegetated SE-facing alpine colluvium, 
Horsfeld Creek valley, 1128 m, 62°2' N 141°11' W, 
Parker & Gracz 6913, 13 August 1996. Wrangell 
Mountains: scattered in loose gravel on steep, SE- 
facing slope, NE slope of Mt. Drum, 1433 m, 62°8.83' 
N 144''30.18' W, C Roland 94-051, 11 June 1994. 

This amphiberingean arctic-alpine fescue is rare in 
Alaska (G4 S2S3) where it grows in dry, rocky situations in 
the subalpine and alpine zones of northern and central 
Alaska. In the Yukon it is known from numerous stations in 
the arctic but only a single locality in the southwest comer 
of the Territory (Cody 1996). The stations cited above 
extend its range southeast 310 km into the Wrangell 
Mountains, 320 km into the Mentasta Mountains and 
450 km into the Nutzotin Moutains from collections in the 
Fairbanks Quad (Wood River Buttes, 64°28.35' N 
148°05.97' W, Du^ et at. 95-80, 95-68, 95-26 & 95-57, 
16 June 1995 (ALA)). These collections also connect the 
range to the northwest in Alaska with the range 155 km to 
the east in the Yukon Territory (Cody 1996). Map 28. 

Festuca minutiflora Rydb., Small-Flower Fescue — 
Chugach Mountains: scattered in dry mineral soil 
of gravel slope, alpine valley near Hanagita Peak, 
1 186 m, 6r4.9r N 143°38.86' W, M. Cook 96418, 
7 July 1996 (ALA). 

This North American cordilleran fescue is new to the 
flora of Alaska. The main range of this species is in the 
Rocky Mountains south to California and New Mexico. It 
is rare in Alaska (G4 SI) and in the Yukon Territory 
(Douglas et al. 1981 ). The collection cited above is 445 km 
west of collections in the Yukon Territory in Kluane 
National Park (Cody 1996). Map 29. 

Festuca richardsonii Hooker (F. rubra L. ssp. 
richardsonii (Hooker) Hulten, Festuca rubra ssp. 
arctica (Hack.) Govor.), Richardson's Fescue — 



Nutzotin Mountains: scattered in mesic graminoid 
stringers between closed low birch willow scrub, 
Beaver Lake, 1341 m, 62^^2.61' N 14r48.39' W, 
Cook & Roland 94197B, 29 June 1994; scattered on 
exposed rubble slope, Wiki Creek, 1411 m, 
61°54.46' N 141°10.68' W, C. Roland 96-074, 
8 June 1996. Wrangell Mountains: gravel bar, 
Nabesna River, 817 m, 62°18.96' N 142°54.07' W, 
C. Roland 96-195, 20 June 1996; occasional in sand 
and gravel on river terrace, Jacksina River, 759 m, 
62°21.5r N 142^52.87' W, C. Roland 96-202, 
21 June 1996; scattered in Rhacomitrium on gravel 
bar across from Black Mountain, 945 m, 62°19.02' N 
143M7.44' W, M. Cook 96720, 8 August 1996; 
sandy area on the edge of alder thicket on floodplain. 
Mud Lake, 1001 m, 62°13.52' N 143°45.28' W, C. 
Roland 96-893, 9 August 1996. 

This species is circumpolar with an arctic-alpine distri- 
bution. The specimens cited above connect its range 
196 km to the northwest in the Alaska Range (Aiken and 
Darby shire 1990) with its range 70 km to the east in the 
Yukon Territory (Cody 1996). Map 30. 

Festuca saximontana Rydberg, Rocky Mountain 
Fescue — Chugach Mountains: open sandy White 
Spruce forest. Tana Dunes, 457 m, 6r4.04' N 
142°45.67' W, M Du^ 91040, 6 July 1991; common 
on margins of lower Tana Dunes, 488 m, 61°6.19' N 
142°55.03' W, M. Cook 94199, C. Roland 94-177A, 1 
July 1994; occasional clumps in sand of open tall 
alder scrub, West Fork Tana River, 448 m, 60°52.95' 
N 142°48.33' W, M. Cook 96565, 1 July 1996; scat- 
tered among alder-willow patches on moraine, alpine 
basin of 12-mile Creek, 1271 m, 60°48.85' N 
142^33.52' W, M. Cook 96631, 1 July 1996 (ALA). 

This fescue occurs across boreal North America and is 
found as far south as Arizona and New Mexico. Our collec- 
tions extend its range 338 km south into the Chugach 
Mountains from a station in the Alaska Range (Hulten 
1968) and connect the range 158 km to the east in the 
Yukon Territory (Cody 1996). Map 31. 

Glyceria pulchella (Nash) K. Schum., MacKenzie 
Valley Mannagrass — Chugach Mountains: mar- 
gin of meadow on river terrace wetlands. Tana River 
flats, 351 m, 6ri2.35' N 142^51.87' W, Dujfy & 
Barnes 96-188, 8 August 1996. 

The specimen cited above extends the range of this 
North American boreal-montane species 337 km south into 
the Chugach Mountains from the Tanana Valley (Hulten 
1968) and connects the range 526 km to the east in the 
Yukon Territory near Carcross (Cody 1996). Map 32. 

Phippsia algida (Soland.) R. Br., Snow Grass — ^ 
Chugach Mountains: scattered in wet gravel, W. Jl 
Fork Goat Creek, 1487 m, 60°59.8' N 142°11.8' W, 
M. Cook 96656, 31 July 1996; moist seepage from 
snowmek. Goat Creek, 1487 m, 60°59.5' N 142°!' 
W, Batten & Barker 96-298, 29 July 1996 (ALA); 
scattered in frost polygons, Verde Ridge, 1554 m, 
61°14.03' N 143°28.52' W, Roland & D'Auria 96- 



2002 



Cook and Roland: Notable Vascular Plants from Alaska 



215 




29. Festuca minutiflora 




30. Festuca richardsonii 




31. Festuca saximontana 



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32. Glyceria pulchella 



216 



The Canadian Field-Naturalist 



Vol. 116 



488, 11 July 1996; occasional in moist calcareous 
gravel, Granite Creek, 1829 m, 61°0.22' N 
14r51.r W, C. Roland 96-736, 28 July 1996. 
Wrangell Mountains: scattered in wet silt, Mud 
Lake, 1001 m, 62°13.52' N 143°45.28' W, C. 
Roland 96-883, 9 August 1996; rare in silt on river 
bar. Copper River, 847 m, 62°28.75' N 143°38.3' 
W, C. Roland 96-934, 12 August 1996. Mentasta 
Mountains: saturated mud seep, headwaters of Lost 
Creek, 62°36.25' N 143°12.08' W, M. Cook 3156, 8 
July 1998. NuTZOTiN Mountains: alkali lake bed, 
Solo Flats, 1335 m, 61°31.55' N 141°25.3' W, M. 
Duffy 92157, 1 July 1992. St. Elias Mountains: 
common in wet seep, 2073 m, 60°57.74' N 
14ri7.33' W, M. Cook 95226, C. Roland 95-192, 
14 July 1995. Wrangell Mountains: rare in moist 
organic soil. Cone Ridge, 2073 m, 62°8.2r N 
143°18.47' W, M. Cook 94367, C. Roland 94-308, 
25 July 1994; occasional in moss, Chetaslina Ridge, 
1615 m, 61°56.51' N 144°25.93' W, M. Cook 
94470, 15 August 1994; frost scars, VABM Sentinel, 
1829 m, 61°39' N 142°32' W, Batten & Barker 96- 
107, 24 July 1996; moist sand along rivulet, Skolai 
Creek, 1341 m, 6r41.5' N 142°23' W, Batten & 
Barker 96-126, 25 July 1996; common in mossy soil 
in seep, Hasen Creek, 1835 m, 61°34.06' N 
142°18.33' W, C. Roland 96-637, 23 July 1996; 
locally abundant in saturated soils, Grotto Creek, 
1661 m, 6r30.56' N 142°24.79' W, C. Roland 96- 
657, 23 July 1996; Lakes Plateau, 1890 m, 62°4.4' N 
143°23.5' W, Leggett & Potkin 95-138, 95-108, 28 
July 1995; abundant in streambed, Jaegar Mesa, 
1893 m, 62°15.9' N 143°1.24' W, C. Roland 95-106, 
30 June 1995; common in seep. Cooper Mountain, 
1942 m, 62^17.16' N 142°31.44' W, C. Roland 94- 
281,24 My 1994. 

This circumpolar arctic-alpine tundra grass is rare in the 
Yukon Territory (Douglas et al. 1981). It had been collect- 
ed in the Wrangell Mountains at Chitistone Pass (Scott 
1968). The collections cited above extend its range 127 km 
into the Chugach Mountains and 135 km into the western 
Wrangell Mountains. Map 33. 

Poa hispidula Vasey (P. macrocalyx Trautv. & C.A. 
Mey.), Large Glume Blue Grass — Chugach 
Mountains: common in clumps on boulder strewn 
area on lower Tana Dunes, 488 m, 61°6.19' N 
142°55.03' W, M. Cook 94201, 94202, C. Roland 
94-177C, 1 July 1994; scattered in saturated, mossy 
fen, vie. Middle Hanagita Lake, 744 m, 61°1 1.58' N 
143°30.49' W, Roland & D'Auria 96-498, 12 July 
1996. Wrangell Mountains: mesic graminoid- 
dwarf scrub tundra, Black Mountain, 1481 m, 
62°20.85' N 143°44.9' W, M. Cook 95/60, 3 July 
1995. 

The specimens cited above represent the northern limit 
of the range of this grass. It is an Alaska-Yukon endemic, 
known from numerous localities along the southern coast 
and the Aleutian Islands. These collections extend its range 
289 km east into the Chugach Mountains and 264 km 



northeast into the Wrangell Mountains from a collection in 
the Anchorage Quad (Ft. Richardson, 61°15.45' N 
149°39.42' W, Duffy & Tande 1008, 2 August 1994 
(ALA)) and connect the range 406 km to the southeast near 
Skagway (Hulten 1968). Map 34. 

Podagrostis aequivalis (Trin.) Scribn. & Merr. 
{Agrostis aequivalvis (Trin.) Trin.), Northern 
Bentgrass — Wrangell Mountains: rare on flood- 
plain of the Dadina River, 393 m, 6r51.49' N 
145°0.72' W, C. Roland 96-308, 1 July 1996. 

The Wrangell Mountains locality represents the northern 
limit of the range of this species, which occurs along the 
west coast of North America, from Oregon to the south 
coast of Alaska. This locality extends the range of this 
species 133 km north into the Wrangell Mountains from a 
station near Cordova (Hulten 1968). Map 35. 

Puccinellia des champ sioides Sorens., Polar Alkalai 
Grass — Wrangell Mountains: near shoreline in 
wet clay, Mt. Drum warm spring, 897 m, 62°4.83' N 
145*^0.55' W, C.R. Meyers 84-90, 18 July 1984; 
sparsely vegetated berm surrounding Lower Klawasi 
Mud volcano, 62°3.52' N 145°13.29' W, M. Cook 
91203, 91204, 21 August 1991. 

This halophytic grass is found in disjunct stations across 
northern North America and western Greenland. The Mt. 
Drum localities represent its western range Umit. These sta- 
tions are 358 km west of a station in the Yukon Territory 
(Cody 1996) where it is considered rare (Douglas et al. 
1981). Map 36. 

Puccinellia interior Sorens., Inland Alkalai Grass — 
NuTZOTiN Mountains: infrequent in saturated moss 
and standing water in graminoid dominated marsh 
adjacent to Ptarmigan Lake airstrip, 1052 m, 61°52.0r 
N 14n0.28' W, M. Cook 3198A, 17 July 1998. 

This species is endemic to Alaska and the Yukon where 
it occurs in moist meadows. It is known from east and 
south-central Alaska and the southern half of Yukon 
Territory, where it is considered rare (Douglas et al. 1981). 
It was collected in the vicinity of the Park at Copper Center 
{C.W. Heideman 2, 1908 (US); J.P. Anderson 2734, 1935 
(H)) and at Chitina {J.P. Anderson 2028, 1 July 1935 (H)) 
(Hulten 1941-1950). Our collection connects the 
distribution from a point 76 km to the east in the Yukon 
Territory (Cody 1996) with those cited above, approxi- 
mately 222 km to the west. Map 37. 

Trisetum sibiricum Rupr. ssp. litorale (Rupr.) 
Roshev., Siberian Oatgrass — Nutzotin 
Mountains: S-facing scree slope, 2.4 km west of 
Ptarmigan Creek, 1494 m, 6r32.89' N 14r3.28' W, 
M. Cook 92482, 9 August 1992; steep S-facing rhyo- 
lite scree knob, west side of Wiki Creek, 1524 m, 
6^54.77' N 14ril.05' W, M. Cook 3190, 15 July 
1998 (ALA). 

This amphiberingean species occurs throughout northern 
Asia, but in North America is restricted to Alaska and a few 
sites in westernmost Yukon Territory where it is considered 
very rare (Douglas et al. 1981). This grass is also consid- 
ered rare in Alaska (G5T4Q 82). Our collections extend its 
range 515 km south into the Nutzotin Mountains and repre- 
sent the southern limit of its distribution in North America. 
Map 38. 



2002 



Cook and Roland: Notable Vascular Plants from Alaska 



217 




33. Phippsia algida 




34. Poa hispidula 




35. Podagrostis aequivalis 




36. Puccinellia deschampsioides 



218 



The Canadian Field-Naturalist 



Vol. 116 



Vahlodea atropurpurea (Wahlenb.) E. Fries, 
Mountain Hairgrass — Gulf of Alaska: mesic tall 
mixed forb herbaceous meadow, Amphitheater 
Knob, 427 m, 59°57' N 139°46' W, M. Cook 87-88, 
19 August 1987; mixed herbaceous meadow, Kan- 
Hills, 549 m, 60°8.66' N 141°16.44' W, K. Beck 87- 
7, 12 August 1987; heather alpine tundra, Floral 
Pass, 792 m, 59°57.91' N 139°57.76' W, M. Cook 
87-100, 21 August 1987; Samovar Hills, 518 m, 
60°9.3' N 140°38' W, M. Cook 87-105, 23 August 
1987. Chugach Mountains: moist soil in Betula 
glandulosa thicket, Martin Creek, 60°56' N 142°23' 
W, Batten & Barker 96-260, 28 July 1996; scattered 
in tall forb herbaceous meadow. Granite Creek, 1344 
m, 60°46.44' n 142°5.12' W, C. Roland 96-805b, 
96-804b, M. Cook 96224, 29 July 1996; common in 
Empetrum heath, 12-Mile Creek, 988 m, 60°50.51' 
N 142°23.02' W, C. Roland 96-764, 29 July 1996; 
common in subarctic lowland sedge wet meadow, 
upper Tebay Lake, 579 m, 61°11' N 144<'24' W, 
Parker & Gracz 6769, 6778, 1 August 1996. 
Wrangell Mountains: patchy in mesic low willow 
forb herbaceous vegetation, Lakina Glacier, 1219 m, 
6r33.64' N 143°19.0r W, M. Cook 96540, C. 
Roland 96-700, 25 July 1996; patchy along shore of 
lake in Empetrum heath, Cheshnina Plateau, 1399 m, 
61°48.04' N 144°6.27' W, M. Cook 96717, 8 August 
1996. St. Elias Mountains: open low shrub, Short 
River Pond, 503 m, 6r5.35' N 14r56.3' W, Parker 
& Duffy 6695, 6 August 1996. 

Vahlodea atropurpurea sens. lat. occurs in meadows and 
thickets throughout the circumpolar north with large gaps 
(Cody 1996). The specimens cited above extend its range 
401 km to the south into the southern St. Elias Mountains 
from a station near Slana (Hulten 1968) and connect the 
western range with a station 385 km to the southeast in the 
Sitka Quad (Freshwater Creek, 58°0.06' N 134°49.57' W, 
C.L. Parker 5251, 14 July 1994 (ALA)). Map 39. 

Cyperaceae 

Carex adelostoma Krecz., Circumpolar Sedge — 
Wrangell Mountains: Fox Farm Lakes, 727 m, 
62°19.98' N 144°50.02' W, M. Potkin 95-008, 26 
July 1995; patchy in mesic graminoid herbaceous 
vegetation, vie. Copper Lake, 911 m, 62°24.53' N 
143°42.68' W, M. Cook 96732, 8 August 1996. 

Carex adelostoma was collected by A. Dutilly, E. 
Lepage and O'Neil near Long Lake on the Nabesna Road 
in 1947 (Lepage 1951; Hulten 1967). Our two collection 
localities are nearby in the upper Copper River watershed. 
These three collections represent the only known localities 
in Alaska and form the western range limit of this species 
in North America. This sedge is rare in Alaska (G4 SI) and 
has an interrupted distribution throughout its range. The 
closest station is 988 km to the northeast at Great Bear 
Lake in the Northwest Territories (Porsild and Cody 1980). 
Map 40. 

Carex alho-nigra Mack., Black-and-White-Scale 
Sedge — Chugach Mountains: S-facing boulder 
slope in unstable gravel scree. Juniper Island, 
1291 m, 60^36.24' N 142^21.69' W, M. Cook 94264, 



11 July 1994. Mentasta Mountains: scattered in 
Arctostaphylos uva-ursi/Festuca altaica meadow on 
S-facing slope, 1097 m, 62°33.3' N 143°9' W, 
Roland & D'Auria 97-042, 26 June 1997; common 
in dry, White Spruce forest at base of bluff, Lost 
Creek, 1097 m, 62°33.3' N 143°9' W, Roland & 
D'Auria 97-077, 26 June 1997. Nutzotin 
Mountains: occasional on lower slopes in Potentilla 
fruticosa/Arctostaphylos uva-ursi community, Carl 
Creek, 1920 m, 62°3.52' N 141^36.27' W, C Roland 
95-043, 20 June 1994; crevices of boulder outcrops 
on S-facing slope, ridge 0.6 km E of Rocker & 
Ptarmigan Creek confluence, 1433 m, 6r54.8' N 
141°2.03' W, M. Cook 94142A, 23 June 1994; 
patchy in talus meadow of S-facing slope. Rock 
Lake, 1119 m, 61M8.7' N 14^16.57' W, 
C. Roland 96-034, 5 June 1996. Wrangell 
Mountains: wet graminoid-dwarf willow-bryoid 
tundra, ridge between Chichokna and Chetaslina 
Rivers, 1615 m, 6r56.51' N 144°25.93' W, M. Cook 
94470A, 15 August 1994. 

This sedge occurs on dry hillsides and talus meadows 
into the subalpine region in eastern Alaska. It is endemic to 
the cordilleran region of western North America, where it 
occurs from New Mexico, Arizona and California north- 
ward into eastern Alaska. It is considered rare in the Yukon 
Territory where it is known from fewer than ten sites 
(Douglas et al. 1981). Our collections extend its range 299 
km southeast into the Nutzotin Mountains and 365 km into 
the Chugach Mountains from a locality in the Alaska 
Range (Black Rapids Roadhouse, 63°32.0' N 145°51.0' W, 
Batten et al. 78-4, 28 June 1978 (ALA)). Map 41. 

Carex buxbaumii Wahlenb., Buxbaum's Sedge — 
Chugach Mountains: sedge meadow. Upper Tebay 
Lake, 579 m, 61°11.5' N 144°23' W, L. A. & E. G. 
Viereck 11093, 15 July 1996; rare in sedge herb 
meadow, Tebay Lake, 579 m, 61°11' N 144°24' W, 
Parker & Gracz 6740, 1 August 1996. Nutzotin 
Mountains: patchy in sphagnum bog. Lick Creek, 
914 m, 62°28.89' N 142°7.56' W, M. Cook 95111, 
28 June 1995. Wrangell Mountains: patchy in 
mesic graminoid herbaceous-open low willow scrub 
guUey, vie. of Whitham Lake, 936 m, 62°19.33' N 
142°53.28' W, M. Cook 96200, 6 June 1996; abun- 
dant at edge of pond, Lakina river, 792 m, 61°29.49' 
N 143°25.6' W, C Roland 96-678, 24 July 1996; 
occasional in marsh, Dadina River, 744 m, 61°53.05' 
N 144°44.09' W, M. Cook 96276A, C. Roland 96- 
344, 2 July 1996. 

This sedge is found in swamps and bogs at low eleva- 
tions across boreal regions of the circumpolar north. 
However, it is absent from the Russian Far-East according 
to Hulten (1968). In North America, it occurs as far south 
as Georgia in the east and California in the west. It is of 
limited distribution in the Yukon Territory (Cody 1996). 
The localities cited above extend its range 189 km to the 
south into the Chugach Mountains from a station near 
Mentasta reported by Hulten (1968). Map 42. 

Carex chordorrhiza Ehrh. ex. L. f.. Creeping Sedge 
— Chugach Mountains: sweetgale/graminoid bog. 



2002 



Cook and Roland: Notable Vascular Plants from Alaska 



219 



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40. Carex adelostoma 



220 



The Canadian Field-Naturalist 



Vol. 116 



I 



Tana River flats, 411 m, 6^12.59' N 142°54.47' W, 
Dujfy & Barnes 96-194, 8 August 1996. Wrangell 
Mountains: numerous in quaking bog, Indian Lake, 
655 m, 62°38.r N 144°20.2' W, M. Cook 95347, 
1 August 1995; locally dominant in wet sedge mead- 
ow, vie. Monte Cristo Creek, 1067 m, 62° 14.27' N 
142°55.81' W, C. Roland 96-136, 18 June 1996; 
co-dominant in sedge marsh, Kuskulana River, 
509 m, 6r28.53' N 143''50.77' W, C. Roland 
96-722, 26 My 1996. 

This sedge occurs in bogs and lake margins from the 
lowlands into the subalpine region in Alaska. Its range is 
circumpolar, but in North Ameica it is generally restricted 
to the boreal region and northward, reaching its southern 
range limit in isolated localities in Illinois. Our collections 
form a range connection for this sedge between a station in 
the Yukon Territory 255 km to the east (Cody 1996) and a 
locality in the Anchorage Quad 226 km to the west (Jim 
and Swan Lakes, 61°33.0' N 148°55.0' W, Batten & Reed 
80-115, 1 July 1980 (ALA)). Map 43. 

Carex crawfordii Fern., Crawford's Sedge — 
Chugach Mountains: dry pond meadow, Tana 
River flats, 442 m, 6ri4.45' N 142°57.21' W, Duffy 
& Barnes 96-140, 8 August 1996. 

This sedge occurs across boreal North America and 
reaches its southern range limit in Missouri (Kartesz 1999). 
It is rare in Alaska (G5 S2S3) and in the Yukon Territory 
(Douglas et al. 1981). The closest collection localities to 
the Tana River are a station near Delta Junction (141 km 
north) and a station near Anchorage 331 km to the west 
(Hulten 1968). Map 44. 

Carex ebumea Boott, Bristleleaf Sedge — St. Elias 
Mountains: White Spruce woodland. Clear Stream, 
488 m, 6r5.82' N 14r57.92' W, Dujfy & Barnes 
96-001, 8 August 1996. Wrangell Mountains: 
patchy in sedge meadow. Grizzly Lake, 1000 m, 
62°19.51' N 143°9.24' W, Cook & Allen 3551, 26 
July 2000. 

These collections extend the range of this North 
American boreal-montane sedge 182 km south into the 
Wrangell Mountains from the Alaska Range (Hulten 1968). 
It had been collected near Chitina (Hulten 1968) and at the 
head of the Chitina River {H.M. Laing s.n., 18 June 1925 
(CAN), Hulten 1941-1950, Porsild 1939). It is rare in 
Alaska (G5 S2S3) and in the Yukon Territory (Douglas et 
al. 1981). Map 45. 

Carex filifolia Nutt., Thread-Leaf sedge — Chugach 
Mountains: steep, SE-facing bluff, confluence of 
Copper and Chitina Rivers, 152 m, 61°31.55' N 
144°24.85' W, C. Roland 94-074B, 12 June 1994; 
steep SW-facing scree slope. Goat Creek, 1487 m, 
60°59.5' N 142°r W, Batten & Barker 96-187B, 
96-224, 27 & 28 July 1996. Mentasta Mountains: 
rare in Arctostaphylos uva-ursi mats on S-facing 
slope. Devil's Mountain, 942 m, 62*'24.95' N 
142^54.86' W, C. Roland 96-231, 22 June 1996. 
Nutzotin Mountains: abundant on steep, S-facing 
bluff, White River between Cub and Traver Creeks, 
1219 m, 6r44.3' N 141°9.5' W, C. Roland 95-078, 
21 June 1995; occasional in well drained areas on 



limestone, Mt. Natazhat, 1716 m, 6r35.38' N 
14ri.83' W, C. Roland 95-060, 19 June 1995. 
St. Elias Mountains: dry sandy mineral soil, vie. 
Walsh Glacier, 1951 m, 60°53.94' N 14r6.75' W, 
M. Cook 95222, 15 July 1995. 

Carex filifolia occurs throughout the western half of 
North America, reaching its eastern limit in Manitoba, 
Canada and its southern limit in New Mexico. It occurs on 
xeric slopes and prairies, and only reaches into eastern 
Alaska, v/here Hulten reported it from two localities, one 
near Chitina (D.F. Murray, B.A. Murray and A. P. 
Khokhryakov s.n., 61°30.0' N 144°25.0' W, 8 July 1981 
(ALA)) and another in the Tanana River valley east of 
Fairbanks. Our collections extend its range into the Nutzotin, 
Chugach, St. Ehas and Wrangell Mountains. Map 46. 

Carex holostoma Drej., Arctic Marsh Sedge — 
Nutzotin Mountains: occasional in open low 
mixed shrub sedge tundra bog, Horsfeld Creek, 1097 
m, 62°1' N 14ril' W, Parker & Gracz 6857 A, 10 
August 1996. 

The specimen cited above extends the range of this 
sedge 1 34 km east into the Nutzotin Mountains from a col- 
lection made by A. Dutilly, E. Lepage and O'Neil near 
Long Lake on the Nabesna Road in 1947 (Lepage 1951; 
Hulten 1967). These collections document the southern 
limit of its range, which is 833 km south of collections on 
the arctic coast of Alaska (Mt. Michelson Quad, Kavik R., 
69°41.0' N 146°52.0' W, M. Emers s.n., 2 July 1992 
(ALA)). This sedge is rare in Alaska (G3G4 S2) and in the 
Yukon Territory (Douglas et al. 1981). Map 47. 

Carex interior Bailey, Inland Sedge — Chugach 
Mountains: occasional in open mossy areas. West 
Fork Tana River, 448 m, 60°52.95' N 142°48.33' W, 
M. Cook 96567, 1 July 1996 (ALA). 

This sedge is disjunct from the temperate zone and is 
rare in Alaska (G5 S 1 ) and in the Yukon Territory (Douglas 
et al. 1981). The specimen cited above extends its range 
288 km east into the Chugach Mountains from a station in 
the Anchorage Quad (Baxter Bog, 6ri3.0' N 149°54.0' 
W, E. F. Layser3275, July 1984 (ALA)). Map 48. 

Carex krausei Boeck., Krause's Sedge — Chugach 
Mountains: wet sedge meadow on low terrace, 
Kiagna River, 762 m, 60°58.33' N 142°20' W, 
Batten & Barker 96-239, 28 July 1996. Mentasta 
Mountains: landslide on W end of Soda Lake, 1 173 
m, 62°32.36' N 142°53.98' W, M. Cook 94306B, 21 
July 1994; occasional in wet sand in openings of 
Populus trichocarpa/Salix alaxensis woodland, 
Totschunda Creek, 725 m, 62°26.94' N 42°40.8' W, 
M. Cook 96254, 6 June 1996. Nutzotin Mountains: 
scattered under Salix pulchra, upper Flat Creek, 
1323 m, 61°58.97' N 141°41.25' W, M. Cook 95036, 
17 June 1995; St. Elias Mountains: open low wil- 
low scrub. Clear Stream, 488 m, 6r5.82' N 
14r57.92' W, Duffy & Barnes 96-029, 96-030, 96- 
031, 8 August 1996. Wrangell Mountains: few in 
bare gravel patches, vie. Boulder Creek, 1036 m, 
62°31.27' N 144''11.26' W, M. Cook 95359, 2 
August 1995; occasional in moist silt of abandoned 
river channel, vie. Black Mountain, 945 m, 






2002 



Cook and Roland: Notable Vascular Plants from Alaska 



221 




41. Carex albo-nigra 




42. Carex buxbaumii 







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43. Carex chordorrhiza 




44. Carex crawfordii 



222 



The Canadian Field-Naturalist 



Vol. 116 




45. Carex eburnea 




46. Carex filifolia 




47. Carex holostoma 



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2002 



Cook and Roland: Notable Vascular Plants from Alaska 



223 




49. Car ex krausei 




50. Carex lasiocarpa ssp. americana 




51. Carex laxa 



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52. Carex lenticularis var. Jr;//V/ 



224 



The Canadian Field-Naturalist 



Vol. 116 



62°19.02' N 143°47.44' W, C. Roland 96-901, 10 
August 1996; patchy in mudflats between gravel bar 
and spruce woodland, Copper River at outflow creek 
from Lake 2990, 884 m, 62°24.38' N 143°42.28' W, 
M. Cook 96724, 8 August 1996. 

This circumpolar arctic-alpine sedge had been collected 
at the head of the Chitna River (H.M. Laing 21, 1925 
(CAN) (Hulten 1941-1950; Porsild 1939)). The collections 
cited above extend its range into the Wrangell, Mentasta 
and Nutzotin Mountains and connect the distribution 
243 km to the north in the Alaska Range (Caster Glacier, 
63°24.0' N 145°43.0' W, Palmer & Porsild 467, 27 June 
1926 (ALA)). Map 49. 

Carex lasiocarpa Ehrh. ssp. americana (Fern.) Hult., 
Wooly-Fruit Sedge — Chugach Mountains: river 
terraced wetlands surrounded by White Spruce for- 
est. Tana River flats, 442 m, 61°14.45' N 142°57.21' 
W, Dujfy & Barnes 96-118, Parker & Gracz 6788, 8 
August 1996. Wrangell Mountains: dense swards 
around small pond, Chokosna Lake, 619 m, 
61°27.45' N 143°49' W, C. Roland 94-330, 10 
August 1994; abundant in wet sedge meadow, vie. 
Dadina River, 744 m, 6r53.05' N 144°44.09' W, C. 
Roland 96-339, 2 July 1996; dominant in pond, 
Kuskulana River, 509 m, 6^28.53' N 143°50.77' W, 
C. Roland 96-720, 96-723, 26 July 1996. 

This North American boreal-montane species is rare in 
the Yukon Territory (Douglas et al. 1981). The specimens 
cited above extend its range east 231 km into the Copper 
and Chitina River basins from a collection in the 
Anchorage Quad (Otter Lake, 6ri7.53' N 149°44.17' W, 
Dujfy & Tande 1017, 3 Aug 1994 (ALA)). Map 50. 

Carex laxa Wahlenb., Weak Sedge — Nutzotin 
Mountains: scattered in marsh. Lick Creek, 914 m, 
62°28.89' N 142°7.56' W, Cook & Beck 95112A, 29 
June 1995. Wrangell Mountains: graminoid 
meadow, confluence of Monte Cristo Creek and 
Nabesna River, 1067 m, 62°14.27' N 142°55.81' W, 
C. Roland 96-134, 18 June 1996. 

The collections of this boreal-montane sedge cited 
above are the only verified stations in Alaska. David F. 
Murray reviewed the collection from Mile 172-174 on the 
Richardson Highway {J. P. Anderson 2712B (S), Hulten 
1941-1950) but could not confirm the original determina- 
tion from the material (personal communication with D. F. 
Murray, 1 1 July, 2000). The collections cited above extend 
its range 368 km to the west from the Yukon Territory. 
This sedge is rare in Alaska (G4 SI) and in the Yukon 
Territory (Douglas et al. 1981). Map 51. 

Carex lenticularis Michx. var. dolia (M.E. Jones) 
L.A. Standley. comb. nov. (C. enanderi Hult.), 
Lakeshore Sedge — Chugach Mountains: flooded 
silt bar of tributary stream, Martin Creek, 1097 m, 
60°56' N 142°23' W, Batten & Barker 96-266, 28 
July 1996; scattered in snowmeit rivulet, west fork 
of 12-mile Creek, 1326 m, 60^50.21' N 142°30.85' 
W, C. Roland 96-787, 29 July 1996; marsh on allu- 
vial fan. Granite Creek, 823 m, 60°44' N 142°13' W, 
Batten & Barker 96-343, 30 July 1996; moist mead- 
ow, west fork of 12-miie Creek, 1326 m, 60^50.21' 



N 142°30.85' W, M. Cook 96593, 1 July 1996 
(ALA). 

This North American cordilleran sedge is rare in Alaska 
(G5T3Q S3). The specimens cited above extend its range 
188 km to the east of a collection in the Valdez Quad 
(Thompson Pass, 61°08.0' N 145°45.0' W, C.L. Parker 
2451, 23 July 1990 (ALA)) and connect the range 423 km 
to the southeast near Skagway (Hulten 1968). Map 52. 

Carex leptalea Wahlenb., Bristly-stalked Sedge — 
Chugach Mountains: sweetgale-graminoid bog. 
Tana River, 411 m, 61°12.59' N 142°54.47' W, 
Dujfy & Barnes 96-196, 8 August 1996; scattered to 
common in small patch of wet sedge herb tundra, 
Tana River, 335 m, 61°ir N 142°51' W, Parker & 
Gracz 6820, 9 August 1996; common in elevated 
mossy hummocks in Sphagnum bog, Hanagita River, 
762 m, 61°11.07' N 143°25.8r W, Roland & 
D'Auria 96-493A, 12 July 1996; common in elevat- 
ed, somewhat drier margin of wet graminoid mead- 
ow, Nerelna Creek, 884 m, 61°25.3' N 144°18.61' 
W, Roland & D'Auria 96-424, 9 July 1996. Gulf of 
Alaska Basin: horsetail bog, Manby Beach, 
59°47.52' N 140°56.51' W, M. Cook 87-117, 20 
August 1987. Nutzotin Mountains: scattered in 
marsh, Lick Creek, 914 m, 62°28.89' N 142°7.56' 
W, M. Cook 95113, 29 June 1995. Wrangell 
Mountains: clumped in Sphagnum at margin of 
pond, Lakina River, 792 m, 61°29.49' N 143°25.6' 
W, M. Cook 96522, 1 July 1996; elevated hummocks 
in wet sedge meadow, Dadina River, 744 m, 
6r53.05' N 144°44.09' W, C. Roland 96-341, 
2 July 1996. 

This sedge is North American with a boreal-montane dis- 
tribution. The localities cited above extend its range south 
220 km into the Nutzotin Mountains, 266 km into the 
Wrangell Mountains and 296 km into the Chugach -i 
Mountains from a station in the Alaska Range (Hulten 1968). f I 
The collection from the Malaspina Forelands extends the 
range 433 east of a station on the Kenai Peninsual (Hulten 
1968). Map 53. 

Carex nardina E. Fries, Nard Sedge — Chugach 
Mountains: rock outcrops, Chakina River, 1646 m, mi 
61°5.3r N 143°0.53' W, M. Cook 94213, 8 July II 
1994; common in Dryas-stdgQ tundra. Juniper 
Island, 1291 m, 60°36.24' N 142°21.69' W, M. Cook 
94271, 12 July 1994; SW-facing rock crevice. 
Juniper Island, 1346 m, 60°36.17' N 142°14.96' W, 
Cook & Roland 94-205, 94250A, 10 July 1994; occa- 
sional in marble rubble. Canyon Creek, 1463 m, 
6r24.35' N 144°20.8' W, Roland & D'Auria 96- 
437, 9 July 1996. Mentasta Mountains: scattered 
in tundra, Totschunda Creek, 1280 m, 62°27.63' N 
142° 12.44' W, C. Roland 96-277, 24 June 1996; dry 
tundra on S-facing slope, Devils's Mountain, 
1530 m, 62°25.62' N 142°53.93' W, C. Roland 96- 
360C, 3 July 1996; common on limestone outcrops. 
Trail Creek, 1615 m, 62°36.05' N 143°17.63' W, C. 
Roland 95-005, 6 June 1995. Nutzotin Mountains: 
limestone gravel, Baultoff Creek, 1707 m, 62°9.13' 



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Cook and Roland: Notable Vascular Plants from Alaska 



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53. Carex leptalea 




54. Carex nardina 




55. Carex nigricans 




56. Carex obtusata 



226 



The Canadian Field-Naturalist 



Vol. 116 



N 141°14.51' W, C Roland 94-164, 28 June 1994; 
limestone ridge, Lime Butte, 1554 m, 6r21.56' N 
142°26.43' W, Cook & Roland 94-232B, 94298B, 14 
July 1994. St. Elias Mountains: occasional on 
limestone gravel, Mt. Natazhat, 1716 m, 61°35.38' 
N 14ri.83' W, M. Cook 95069, 19 June 1995; com- 
mon in disturbed marble rubble, Blondie Ridge, 
1951 m, 60°53.94' N 141°6.75' W, C Roland 95- 
152, 11 July 1995. Wrangell Mountains: gravelly 
limestone, Chitistone Mountain, 1219 m, 6r28' N 
142°33' W, Batten & Barker 96-141, 26 July 1996; 
scattered in gravelly tundra, Cheshnina Plateau, 1399 
m, 61°48.04' N 144°6.27' W, C. Roland 96-854, 6 
August 1996. 

This sedge is amphi-atlantic with an arctic-alpine distri- 
bution. It was collected in the Wrangell Mountains at 
Bonanza Ridge (Nordell and Schmitt 1978) and Guerin 
Glacier (6r37.0' N UrSO.O' W, D.F. Murray 2066, 3 
August 1968 (ALA)). The localities cited above extend its 
range 118 km east from a station in the Alaska Range 
(Hulten 1968) and 102 km south into the Chugach 
Mountains from the Bonanza Ridge collection. Map 54. 

Carex nigricans C.A. Mey, Blackish sedge — 
Chugach Mountains: alpine herbaceous meadow, 
Karr Hills, 549 m, 60°8.66' N 141°16.44' W, K.A. 
Beck 87-11, 12 August 1987; occasional in 
Empetrum heath, 12-mile Creek, 988 m, 60°50.51' N 
142°23.02' W, C. Roland 96-766A, 29 July 1996; 
scattered in snowmelt rivulet, 12-mile Creek, 
1326 m, 60°50.21' N 142°30.85' W, C Roland 96- 
788, 29 July 1996; scattered in lush forb-herbaceous 
meadow. Granite River, 1344 m, 60M6.44' N 
142°5.12' W, C. Roland 96-800, 29 July 1996; occa- 
sional in wet moss, vie. Spirit Mountain, 1006 m, 
6ri6.84' N 144°29.86' W, M. Cook 96433, 1 July 
1996; occasional in Cassiope/Luetkea heath of alpine 
basin, vie. 12-mile Creek, 1271 m, 60°48.85' N 
142°33.52' W, M. Cook 96626, 1 July 1996. Gulf of 
Alaska: Cassiope alpine tundra. Samovar Hills, 549 
m, 60°8.ir N 140°39.55' W, M. Cook 87-102, 23 
August 1987; gravelly area next to wet equisetum 
meadow, Guyot Glacier, 183 m, 60°3.6' N 141°18' 
W, K.A. Beck 92545, 22 August 1992. Wrangell 
Mountains: ridge 0.6 km S of Nadina Glacier, 
1768 m, 62°2.85' N 144^41' W, M. Cook 94424A, 
30 August 1994. 

The northern limit of the range of this North American 
cordilleran sedge is documented by the collection cited 
above from the Wrangell Mountains. These collections 
extend its range 1 16 km to the northeast into the Wrangell 
Mountains, 203 km east into the Chugach Mountains and 
291 km southeast into the southern St. Elias Mountains 
from a station near Thompson Pass (Hulten 1968) and con- 
nect the range 298 km to the southeast near Skagway 
(Hulten 1968). Map 55. 

Carex obtusata Liljeb., Blunt Sedge — Chugach 
Mountains: rubbly area of SW-facing slope underlain 
by ultramafic rock, Chakina River, 1768 m, 6r6.19' 
N 142°55.03' W, C. Roland 94-185B, 8 July 1994; dry 



slopes, Goat Creek, 1487 m, 60°59.5' N 142°1' W, 
Batten & Barker 96-213, 27 July 1996; dry soil on 
exposed bench, Goat Creek, 1487 m, 60°59.5' N 
142°1' W, Batten & Barker 96-316, 29 July 1996 
(ALA). Nutzotin Mountains: scattered in turfy, 
Kobresia dominated community on knoll. Rock Lake, 
1119 m, 6r48.7' N 14ri6.57' W, C Roland 96-033, 
5 June 1996. Wrangell Mountains: Drya^/dwarf 
willow tundra community. Moose Point, 1052 m, 
62°31.61' N 144°12.38' W, C. Roland 94-021, 1 June 
1994; occasional on dry hillside, Nabesna River 
bluff, 1106 m, 62^5.05' N 142°54.68' W, C. Roland 
96-157, 19 June 1996; few in bare mineral soil on 
well drained slope, Nabesna River bluff, 1106 m, 
62°15.05' N 142^54.68' W, M. Cook 96192, 6 June 
1996. 

This species is amphiberingean with an arctic-alpine 
distribution. The specimens cited above extend its range 
south 169 km into the Wrangell Mountains, 244 km into 
the Nutoztin Mountains and 315 km into the Chugach 
Mountains from a collection in the Alaska Range (Mt. 
Hayes Quad: 63°43.0' N 144°23.0' W, Batten & Juday 
85-207, 12 July 1985 (ALA)) and connect the range 
126 km to the east in the Yukon Territory (Cody 1996). 
Map 56. 

Carex parry ana Dew., Parry's Sedge — St. Elias 
Mountains: open low willow scrub, Clear Stream, 
terminus of Barnard Glacier, 488 m, 6r5.82' N 
14^57.92' W, Duffy & Barnes 96-012, 8 August 
1996. 

This North American boreal-montane sedge is rare in 
Alaska (G4 SI). The specimen cited above extends the 
range in the Chugach Mountains 348 km to the east from a 
collection in the Anchorage Quad (Eklutna, 61°28.0' N 
149°22.0' W, Dutilly et al. 20709, 1 July 1947 (ALA)) and 
is 162 km west of collections in the Yukon Territory (Cody 
1996). Map 57. 

Carex pauciflora Lightf., Star Sedge — Chugach 
Mountains: boggy area in open tall willow alder 
scrub. Falls Creek, 610 m, 6ri0.58' N 144°24.75' 
W, M. Duffy 91103, 12 August 1991; ericaceous 
shrub bog. Middle Fork of the Bremner River, 869 m, 
60°55.05' N 143°43.86' W, Duffy & Barnes 96-093, 
8 August 1996. Gulf of Alaska: ericaceous 
shrub/graminoid herbaceous bog, Robinson 
Mountains, 427 m, 60°4.8' N 142''12.8' W, M Duffy 
9216, 11 June 1992. 

The collections cited above extend the range of this bore- 
al-montane sedge east 1 1 1 km in the Chugach Mountains 
and 206 km into the Robinson Mountains from a station near 
Cordova (Orca, 60°39.83' N 145°43.00' W, I.L Norberg s.n., 
28 August 1938 (S), Hulten 1941) and connect the range 
227 km to the southeast near Yakutat (Lost River area, 
59°28.0' N 139°40.0' W, C. L. Parker 2611, 24 June 1991 
(ALA)). Map 58. 

Carex petasata Dew., Liddon Sedge — Chugach 
Mountains: locally common in a dry meadow on 
lateral moraine. Granite Creek, 884 m, 60°44.63' N 
142°6.05' W, C. Roland 96-791, 29 July 1996 

(ALA). 



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5 7. Ca rex pa rryana 




58. Carex pauciflora 




59. Carex petasata 




60. Carex petricosa 



228 



The Canadian Field-Naturalist 



Vol. 116 



This North American cordilleran sedge is new to the 
flora of Alaska and considered rare (G5 SI). The collection 
cited above is 262 km west of stations in the southwest 
Yukon near Haines Junction (Cody 1996) and represents 
the western limit of its known range. Map 59. 

Carex petricosa Dewey, Rockdwelling Sedge — 
Chugach Mountains: dry soil on steep SW-facing 
slope, Goat Creek, 1487 m, 60°59.5' N 142°1' W, 
Batten <5: Barker 96-219, 27 July 1996; occasional 
patches in gravel and finer scree, vie. Iron Creek, 
1612 m, 6ri.49' N 141°54.23' W, M. Cook 96580, 1 
July 1996. Mentasta Mountains: densely vegetated 
shoulder amid extensive gravel chutes on SW-facing 
slope, Totschunda Creek, 1280 m, 62°27.63' N 
142°12.44' W, C. Roland 96-296, 24 June 1996. 
Nutzotin Mountains: subalpine meadow at treeline, 
Carden Hills, 1311 m, 62^8.44' N 141°! 1.55' W, C. 
Roland 94-135, 25 June 1994. St. Elias Mountains: 
common in dry tundra on limestone, ridge between 
Dan & Copper Creeks, 1554 m, 6r21.56' N 
142°26.43' W, C. Roland 94-231B, 14 July 1994; 
occasional in well-drained areas on limestone, Mt. 
Natazhat, 1716 m, 61°35.38' N 14ri.83' W, C. 
Roland 95-059, 95-061, 19 June 1995; in limestone 
scree with stringers of Dryas alaskensis, Mt. 
Natazhat, 1716 m, 6r35.38' N 14ri.83' W, M. Cook 
95078A, 21 June 1995. Wrangell Mountains: occa- 
sional in open, subalpine White Spruce forest and 
meadows on W-facing slope of Hmestone knoll. Fish 
Creek, 1067 m, 62°16.92' N 142°58.99' W, C. Roland 
95-i05, 30 July 1995. 

This North American cordilleran sedge had been collect- 
ed in the St. Elias Mountains (Guerin Glacier, 61°37.42' N 
141°4.38' W, D. F. Murray 2063, 3 August 1968 (ALA)). 
The collections cited above extend its range 91 km south 
into the Chugach Mountains, 111 km north into the 
Nutzotin Mountains and 126 km northwest into the 
Wrangell Mountains. These collections also connect the 
range 203 km to the northwest in the Alaska Range (Mile 
238 Richardson Highway, 63°40.0' N 145°52.0' W, Batten 

6 Dawe 78-123, 4 July 1978 (ALA)) with the range 
134 km to the east in the Yukon Territory (Cody 1996). 
Map 60. 

Carex phaeocephala Piper, Dunhead Sedge — 
Chugach Mountains: scattered in loose, shaly talus 
on S-facing slope, vie. 12-mile Creek, 1335 m, 
60M9.7r N 142°33.34' W, C. Roland 96-815, 
30 July 1996; tufted on slope of moraine in 
Arctostaphylos iiva-ursi patches. Granite River, 
884 m, 60''44.63' N 142°6.05' W, M. Cook 96602, 

7 July 1996; mesic shrub birch-ericaceous. Granite 
Creek, near Ross Green Lake, 701 m, 60°43.83' N 
142°28.78' W, Duffy & Barnes 96-056, 96-057, 

8 August 1996. 

This North American cordilleran sedge is rare in the 
Yukon Territory (Douglas et al. 1981) and known from 
only five localities in Alaska. The collections cited above 
extend its range 104 km east in the Chugach Moutains from 
a collection near Cordova (Schwan Glacier, 60°58.0' N 
145°00.0' W, C Parker 1763, 12 August 1986 (ALA)) and 



connect the range 1 34 km to the east in the Yukon Territory 
(Cody 1996). Map 61. 

Carex praticola Rydb., Meadow Sedge — Chugach 
Mountains: silt of ledges. Juniper Island, 1346 m, 
60°36.17' N 142°14.96' W, Cook & Roland 94-204, 
94254, 10 July 1994; scree on S-facing boulder 
slope, Juniper Island, 1291 m, 60°36.24' N 
142°21.69' W, M. Cook 94260, 11 July 1994; moist 
gravel, Short River Pond, 503 m, 61°5.35' N 
141°56.3' W, Parker & Duffy 6686, 6 August 1996; 
edges of pools in marshy area, Granite River, 823 m, 
60°44' N 142°13' W, Batten & Barker 96-344, 
30 July 1996. 

This sedge is North American with a boreal-montane 
distribution. The localites cited above extend its range 
244 km east in the Chugach Mountains from a station near 
Valdez (Hulten 1968) and connect its range 283 km to the 
east in the Yukon Territory near Haines Junction (Cody 
1996). Map 62. 

Carex stylosa C.A. Mey, Variegated Sedge — 
Chugach Mountains: muddy margin of small kettle 
pond. Granite Creek, 701 m, 60°43.78' N 142°31.4' 
W, Duffy & Barnes 96-066, 8 August 1996; scattered 
in wet meadow, Granite River, 884 m, 60°44.63' N 
142°6.05' W, C. Roland 96-795B, 29 July 1996; 
sedge meadow, Upper Tebay Lake, 579 m, 61° 11.5' 
N 144°23' W, L. A. & E. G. Viereck 11096, 7 July 
1996; mesic sedge herb meadow, Tebay Lake, 
579 m, 6ril' N 144°24' W, Parker & Gracz 6725, 
67 67 A, 1 August 1996; locally common in mossy 
heath, 12-Mile Basin, 1335 m, 60^49.71' N 
142°33.34' W, C Roland 96-830, 30 July 1996; eri- 
caceous shrub bog. Middle Fork of the Bremner 
River, 869 m, 60°55.05' N 143°43.86' W, Duffy^ & 
Barnes 96-091, 96-100, 8 August 1996. Wrangell 
Mountains: common in mud on tundra pond, 
Cheshnina Plateau, 1399 m, 6r48.04' N 144°6.27' 
W, C. Roland 96-857, 6 August 1996. 

This amphiberingean boreal-montane sedge is 
rare in the Yukon Territory (Douglas et al. 1981). 
The collections cited above extend its range 135 
northwest into the Wrangell Mountains and 214 east 
into the Chugach Mountains from a station near 
Worthington Glacier (6ri.08' N 146°34' W, LA. 
Viereck 8475, 1 August 1967 (ALA)). Map 63. 

Carex viridula Michx. (C. oederi Retz. ssp. viridula 
(Mischx.) Hult.), Little Green Sedge — Chugach 
Mountains: rare in wet silt on floodplain, conflu- 
ence of Copper and Bremner Rivers, 152 m, 
60°56.95' N 144M1.79' W, C. Roland 96-610, 
17 July 1996. Gulf of Alaska: wet sedge bog, 
Sudden Stream, 30 m, 59^30.71' N 139°46.44' W, 
M. Cook 8850, 20 July 1992; gravelly area next to 
wet Equisetum meadow, Guyot Glacier, 183 m, 
60°3.6' N 14ri8' W, M. Cook 92552, 19 August 
1992. St. Elias Mountains: wet fen in black spruce 
woodland, Bryson Bar, 1800 m, 61M.57' N 
14r54.47' W, M. Duffy^ 91029, 3 July 1991; mixed 



II 






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Cook and Roland: Notable Vascular Plants from Alaska 



229 



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61. Carex phaeocephala 




62. Carex praticola 




63. Carex stylos a 




64. Carex viridula 



230 



The Canadian Field-Naturalist 



Vol. 116 



graminoid-forb meadow, Clear Stream, 488 m, 
6r5.82' N 14r57.92' W, Dujfy & Barnes 96-034, 
8 August 1996. Wrangell Mountains: mud flats 
around Lake 2870, vie. Tanada Creek, 875 m, 
62°31.38' N 143^28. 12' W, Moran & Roland 95-08, 
28 June 1995. 

This amphiberingean boreal-montane sedge is rare in the 
Yukon Territory (Douglas et al. 1981) and known from few 
localities in Alaska. The collections cited above extend its 
range 190 km south into the Wrangell Mountains from a sta- 
tion in the Alaska Range (Hulten 1968), 337 km east in the 
Chugach Range from a collection in the Anchorage Quad 
(Bulldog Road bogs, 61°13.10' N 149°41.40' W, Dujfy & 
Tande 1065, 5 August 1994 (ALA)) and 103 km to the west 
on the Malaspina Forelands from a station at Yakutat 
(59°29.86' N 139°45.60.' W, Stair & Pennell s.n., 1945 
(PH), Hulten 1941-1950, Stair and Pennell 1947). Map 64. 

Carex williamsii Britt., Williams' Sedge — Nutzotin 
Mountains: wet sedge meadow tundra at margin of 
kettle lake, northwest of Beaver-Horsfeld Creek con- 
fluence, 1097 m, 62°1' N 141°! 1' W, Parker & Gracz 
6844, 10 August 1996. 

This amphiberingean arctic-alpine sedge is rare in the 
Yukon Territory (Douglas et al. 1981). The collecdon cited 
above extends its range 205 km southeast into the Nutzotin 
Mountains from a station in the Alaska Range (Hulten 
1968) and connects the range 155 km to the east in the 
Yukon Territory (Cody 1996). Map 65. 

Eriophorum callitrix Cham., Arctic Cotton-grass — 
Chugach Mountains: moist soil near edge of pond, 
Martin Creek, 1097 m, 60°56' N 142°23' W, Batten 
& Barker 96-246 (ALA), 96-264, 28 July 1996 
(ALA). Wrangell Mountains: Lakes Plateau, 1890 
m, 62°4.4' N 143°23.5' W, M. Potkin 95-100, 30 
July 1995; alpine basin, NW flank of Mt. Drum, 
1615 m, 62°4.5' N 144^45.91' W, C. Meyers 84-23, 
4 July 1984; scattered in wet graminoid meadow, 
Chetaslina Ridge, 1615 m, 6r56.51' N 144°25.93' 
W, C. Roland 94-360, 17 August 1994. 

This circumpolar arcdc-alpine grass was collected in the 
Wrangell Mountains at Chitistone Pass (Scott 1968). The 
localities cited above extend its range 151 km west into the 
western Wrangell Mountains and 87 km south into the 
Chugach Mountains. Map 66. 

Eriophorum viridi-carinatum (Engelm.) Fern., Tassel 
Cotton-grass — Wrangell Mountains: occasional in 
Sphagnum bog, Lakina River, Chitina River Basin, 
792 m, 6r29.49' N 143°25.6' W, C Roland 96-672, 
24 July 1996. 

This North American boreal-montane cotton-grass is 
rare in Alaska (G5 S2) and in the Yukon Territory (Douglas 
et al. 1981). The specimen cited above connects its distri- 
bution 259 km to the east in the Anchorage Quad (Muldoon 
bog, 61° 12.22' N 149°42.97' W, Duffy & Tande 923, 
26 July 1994 (ALA)) with a station 458 km to the southeast 
near Haines (Hulten 1968). Map 67. 

Kobresia sibirica Turcz., Siberian Bog Sedge — 
Nutzotin Mountains: patchy on upper slope of S- 
facing bluff. White River, 1219 m, 6r44.3' N 
14r9.5' W, M. Cook 95085, 21 June 1995. 



This species is circumpolar with an arctic-alpine distri- 
bution. The specimen cited above extends its range 270 km 
to the southeast into the Nutzotin Mountains from a collec- 
tion in the Alaska Range (Bear Creek, 63°37.0' N 
145°50.0' W, Batten & Dawe 78-42, 29 June 1978 (ALA)) 
and connects the range 62 km to the east in the Yukon 
Territory (Cody 1996). Map 68. 

Kobresia simpliciuscula (Wahlenb.) Mack., Simple 

Bog Sedge — Mentasta Mountains: co-dominant 
in hummocky meadow on floodplain. Lost Creek, 
1006 m, 62^32' N 143°9.6' W, Roland & D'Auria 
97-062, 97-073, 26 June 1997. Nutzotin Moun- 
tains: mesic graminoid herbaceous meadow 
between Gold Run and Glacier Creeks, 1493 m, 
62''7.42' N 141°52.73' W, M. Cook 8983, 23 July 
1989; wet sedge meadow tundra at margin of kettle 
lake, Beaver-Horsfeld Creek confluence, 1097 m, 
62°1' N 14ril' W, Parker & Gracz 6843, 6857, 
6900, 10 August 1996; sedge meadow, N of Francis 
Creek, 1234 m, 6r53.78' N 141°8.81' W, M. Du_ffy 
96-273, 8 August 1996. 

The collections cited above of this circumpolar arctic- 
alpine sedge extend its range 136 km south into the 
Mentasta Mountains and 244 southeast into the Nutzotin 
Mountains from the Alaska Range (Hulten 1968) and 
connect its distribution 130 km to the east in the Yukon 
Territory (Cody 1996). Map 69. 

Trichophorum pumilum (M. Vahl.) Schinz. & Thell. 
var. rollandii (Scirpus rollandii Fern.), Rolland's 
Leafless-Bulrush — St. Elias Mountains: wet allu- 
vium at terminus of Barnard Glacier, St. Elias 
Mountains, 488 m, 6r5.82' N 141°57.92' W, Duffy 
& Barnes 96-036, 5 August 1996. 

This boreal montane sedge is new to the state of Alaska. 
The collection cited above represents the western limit of 
its range and is 191 km west of collections in the Yukon 
Territory (Cody 1996). It is rare in Alaska (G3T3T4? SI) 
and in the Yukon Territory (Douglas et al. 1981). Map 70. 

JUNCACEAE 

Juncus filiformis L., Thread Rush — Chugach 
Mountains: abundant in mud. Granite River, 884 m, 
60°44.63' N 142°6.05' W, C Roland 96-790, 29 July 
1996; scattered at margin of subarctic lowland sedge 
wet meadow, upper Tebay Lake, 579 m, 61°ir N 
144^24' W, Parker & Gracz 6738, 1 August 1996. 
St. Elias Mountains: wet sedge herb meadow tun- 
dra. Short River, 503 m, 61°5.35' N 141°56.3' W, 
Parker & Dujfy 6681, 6 August 1996. Wrangell 
Mountains: scattered on sandy beaches and alluvial 
flats, Fox Farm Lake, 727 m, 62°19.98' N 
144°50.02' W, C. Roland 95-260, 25 July 1995; 
occasional in bare clay soil in graminoid meadow, 
SE end Boulder Lake, 1036 m, 62°31.27' N 
144°1 1.26' W, M. Cook 95365, 2 August 1995; com- 
mon along lake shore in open silt. Copper River, 91 1 
m, 62°24.53' N 143°42.68' W, C. Roland 96-917, 
1 1 August 1996. 

This rush is circumpolar with a boreal-montane distribu- 
tion. The collections cited above extend its range southeast 






2002 



Cook and Roland: Notable Vascular Plants from Alaska 



231 




65. Carex williamsii 




66. Eriophorum callitrix 




67. Eriophorum viridi-carinatum 




68. Kobresia sibirica 



232 



The Canadian Field-Naturalist 



Vol. 116 



103 km into the Wrangell Mountains and 330 km into the 
Chugach Mountains from a, station in the Alaska Range 
(Mile 3.3 Denah Highway, 63°40.0' N 145°33.0' W, LA. 
Viereck 8359, 26 July 1967 (ALA)). Map 71. 

Juncus mertensianus Bong., Merten's Rush — 
Chugach Mountains: rocks and saturated silt, 
lower Bremner River, 488 m, 6r0.8r N 144°14.39' 
W, C. Meyers 84-104, 20 July 1984; sand in open 
willow scrub. Granite River, 884 m, 60°44.32' N 
142°4.06' W, M. Cook 96616, 1 July 1996; scattered 
forbs. Short River, 503 m, 61°5.35' N 14r56.3' W, 
Parker & Dujfy 6691, 6 August 1996; scattered in 
moss, 12-Miie Basin, 1326 m, 60^50.21' N 
142°30.85' W, M. Cook 96599, 1 July 1996; along 
stream in forb herbaceous vegetation, Falls Creek, 
655 m, 61°14.17' N 144°28.24' W, L. A. & E. G. 
Viereck 11044, 1 July 1996; tall willow scrub. 
Granitic Creek, 1188 m, 6r4.37' N 142°56.79' W, 
M. Hoffman s.n., 11 August 1986; in floating peat on 
lakeshore, Martin Creek, 1097 m, 60°56' N 142°23' 
W, Batten & Barker 96-257, 28 July 1996; abundant 
in meadow. Canyon Creek, 1035 m, 61°20.17' N 
144°19.38' W, C. Roland 96-549, 15 July 1996; 
common in heath, 12-mile Creek, 988 m, 60°50.5r 
N \M°lZm' W, C. Roland 96-765, 1 July 1996; 
seepage area at margin of subarctic lowland sedge 
wet meadow. Granite Creek, 701 m, 60°43.8r N 
142°30.08' W, Parker & Gracz 6773, 8 August 
1996. St. Elias Mountains: herbaceous meadow, 
Karr Hills, 60°8.66' N 141°16.44' W, K. Beck s.n., 
12 August 1987. Wrangell Mountains: patchy in 
wet mud. Grant Creek, 853 m, 61°18.16' N 
143°51.87' W, Cook & Losso 96334, 1 July 1996; 
moist fine sand along rivulet, Skolai Creek, 1341 m, 
6r41.5' N 142°23' W, Batten & Barker 96-129, 25 
July 1996; patchy in graminoid-forb meadow, 
Cheshnina Plateau, 1399 m, 61°48.04' N 144°6.27' 
W, C. Roland 96-869, 7 August 1996. 

This rush is North American with a cordilleran distribu- 
tion. The specimens cited above extend its range east 
225 km into the Chugach Mountain, 285 km into the south- 
ern St. Elias Mountains and northeast 211 km into the 
Wrangell Mountains from a collection in the Valdez Quad 
(61° 12.0' N 145°47.0' W, L.A. Viereck 8472, 1 August 
1967 (ALA)). These localities also connect the range 
151 km to the east in the Yukon Territory (Cody 1996). 
Map 72. 

LiLIACEAE 

Maianthemum stellatum (L.) Link., Star-flowered 
Solomon's Seal — Nutzotin Mountains: Populus 
balsamifera forest on steep slope, between Sheep 
and Notch Creeks, 488 m, 62° 15.09' N 14r59.17' 
W, D. Morrison 84-41, 19 June 1984. 

This lily is North American with a boreal-montane dis- 
tribution and is rare in Alaska (G5 S2). The specimen cited 
above extends its range 328 km to the east into the Nutzotin 
Mountains from a collection in the Anchorage Quad (Long 
Lake, 61°48.55' N 148° 14.56' W, M. Cook 3114, 2 June 
1998 (ALA)) and 292 km south from collections in the 
Eagle Quad (Mission Creek, 64°48.0' N 141°10.0' W, 



Khokhryakov et al. 6294, 10 July 1981 (ALA)). This locali- 
ty also connects the distribution 170 km to the east in the 
Yukon Territory (Cody 1996). Map 73. 

Salicaceae 

Salix commutata Bebb, Undergreen Willow — 
Chugach Mountains: flooded gravel bar, Martin 
Creek, 1097 m, 60°56' N 142°23' W, Batten & 
Barker 96-262, 28 July 1996; scattered in wet silt with 
Equisetum, Copper/Bremner River confluence, 
152 m, 60°56.95' N 144°41.79' W, C. Roland 96-605, 
17 July 1996. Gulf of Alaska Basin: graminoid 
herbaceous meadow. Alder Stream, 59°44.22' N 
140°22.95' W, M. Cook 87-94, 16 August 1987; 
horsetail bog. Cape Sitkagi, 61 m, 59°47.52' N 
140°56.5r W, M. Cook 87-99, 20 August 1987; wet 
sedge marsh, Esker Stream, 59°54.19' N 139°46.27' 
W, K. Beck s.n., August 1987. 

The specimens cited above of this North American 
cordilleran willow extend its range 148 km to the east into 
the Chugach Mountains from a station on the Copper River 
near the Bremner River (Hulten 1968) and connect the 
range 90 km to the southeast near Yakutat (Harlequin Lake, 
59°24.0' N 139°00.0' W, L.A. Viereck s.n., 7 July 1965 
(ALA)). Map 74. 

Salix rotundifolia Trautv. ssp. dodgea